E N V R ON M E N T 21653 ~~~~~D E P A R-T M E N T ( 6) ~~P A P E R -SPAE.7-. PAPERS ~~~~~~~~~~~PAPER N.75* 4dw ~ TOWARD ENVIRONMENTALLY AND SOCIALLY SUSTAINABLE DEVELOPMENT B.IODIVERSITY SERIES - IMPACT STUDIES Biodiversity Conservation in the Context of Tropical Forest Management Francis E. Putz Kent H. Redford John G. Robinson Robert Fimbel' Geoffrey M. Blate September 2000 The World Bank O THE WORLD BANK ENVIRONMENT DEPARTMENT Biodiversity Conservation in the Context of Tropical Forest Management Francis E. Putz Kent H. Redford John G. Robinson Robert Fimbel Geoffrey M. Blate September 2000 Papers in this series are not formal publications of the World Bank. They are circulated to encourage thought and discussion. The use and citation of this paper should take this into account. The views expressed are those of the authors and should not be attributed to the World Bank. Copies are available from the Environment Department, The World Bank, Room MC-5-126. The International Bank for Reconstruction and Development/THE WORLD BANK 1818 H Street, N.W. Washington, D.C. 20433, U.S.A. Manufactured in the United States of America First printing September 2000 About the authors: Francis E. Putz (fep@botany.ufl.edu) is Professor of Botany at the University of Florida; Kent H. Redford (KHRedford@aol.com) is Director of Biodiversity Analysis and Coordination at WCS; John G. Robinson (jrobinson@wcs.org) is Senior Vice President and Director for International Conservation at WCS; Robert Fimbel (robertf@parks.wa.gov) is Chief Scientist for the Washington State Parks and Recreation Commis- sion; and Geoffrey M. Blate (gblate@botany.ufl.edu) is a Ph.D. candidate at the University of Florida. Substantial background material for this paper was provided by D. Nepstad, M. Guariguata, R. Noss, and C. Peters. We also acknowledge the contributions of E. Bennett, J. Ewel, P. Frumhoff, E. Losos, K. MacKinnnon, A. Moad, G. Platais, A. Plumptre, and R. Plumptre. This document does not represent an endorsement of any particular World Bank policy by the Wildlife Conservation Society. 'WC s The Wildlife Conservation Society (WCS) is dedicated to being the most effective conservation organiza- tion, protecting and promoting a world rich in wildlife and wilderness. WCS manages more than 300 field projects in 53 countries and develops award-winning environmental education programs for schools in the U.S. and abroad. At its celebrated wildlife parks in New York-the Bronx Zoo, the New York Aquarium, and Central Park, Queens and Prospect Park Wildlife Centers-WCS inspires over 4.5 million annual visitors to care about wildlife and wild lands and to participate in their conservation. To leam more about WCS, visit www.wcs.org. Biodiversity Series Impact Studies These "Impact Studies" are a subset of the Biodiversity Series of the World Bank's Environment Department Papers. Within this subset the broader question of what the positive and negative impacts of human activities on biodiversity is addressed. The following studies have been published in this series: 1. Biodiversity Conservation in the Context of Tropical Forest Management 2. Hunting of Wildlife in Tropical Forests-Implicationsfor Biodiversity and Forest Peoples Contents PREFACE V ExEcurlvE SUMMARY VIl Chapter 1 Introduction 1 Chapter 2 Disaggregating "Biodiversity" 3 Chapter 3 Disaggregating "Logging" 7 Logging intensities 7 Log yarding methods 9 Logging and other silvicultural treatments 11 Reducing the impacts of logging and other silvicultural treatments 12 Chapter 4 Impacts of Forest Management on Biodiversity 15 Background 15 Landscape impacts (see Appendix I) 16 Ecosystem impacts (see Appendix II) 19 Community impacts (see Appendix III) 21 Species impacts (see Appendices V and VI) 22 Genetic impacts (see Appendix VII) 25 Chapter 5 Overview of Biodiversity Conservation in Relation to Logging and other Silvicultural Treatments 27 Synthesis 27 Troubling assumptions 29 Conclusions 30 Chapter 6 Recommendations 33 Biodiversity Series - Impact Studies iii Biodiversity Conservation in the Context of Tropical Forest Management APPENDICES I Impacts of Logging (and other silviculture treatments where noted) on the Landscape Component of Biodiversity in Tropical Forests 35 II Impacts of Logging (and other silviculture treatments where noted) on the Ecosystem Component of Biodiversity in Tropical Forests 39 III Impacts of Logging (and other silviculture treatments where noted) on the Community Compo- nent of Biodiversity in Tropical Forests 43 IV Bird responses (density based on inter-site comparisons) to Logging in Tropical Forests, Organized by Feeding Guild 51 V Impacts of Logging (and other silviculture treatments where noted) on the Species Component of Biodiversity in Tropical Forests 53 VI Primate responses (by feeding guild) to Logging in Tropical Forests 57 VII Impacts of Logging (and other silviculture treatments where noted) on the Genetic Component of Biodiversity in Tropical Forests 61 GLOSSARY 63 BIBLIOGRAPHY 67 Boxes 1 Few tropical forests are logged or burned only once 9 2 Enrichment planting 12 3 Is reduced-impact logging (RIL) cheaper? 13 4 The critical role of production forests in maintaining biodiversity in the tropics 17 5 Hunting in logged tropical forests 23 6 Lesser-known species-Marketing challenges, silvicultural impediments, or desirable diversity? 24 Figures 1 Logging intensities (m3/ha) for tropical forests 8 2 Generalized ranges of direct biodiversity impacts of timber yarding methods used for tropical forest logging as a function of level of technological sophistication required 10 3 Some impacts of logging on biodiversity as a function of distribution of logging activities and logging intensity 18 4 Expected effects of a range of forest uses on the components of biodiversity 27 5 Generalized biodiversity impacts plotted against expected short-term financial returns to forest owners or concessionaires 28 6 Generalized biodiversity impacts resulting from different approaches to forest management for timber 29 Tables 1 Components and attributes of tropical forest biodiversity that might be influenced by logging and other silvicultural activities 4 iv Environment Departrnent Papers Preface This paper was prepared as part of the World One of the most important forest uses is logging Bank's Overlay Program and as a contribution or timber harvesting. Logging is the most to the Bank's ongoing Forest Policy review. lucrative forest use and can cause severe direct The Overlays Program, launched by the World and indirect environmental impacts. This Bank in partnership with bilateral donors and analysis evaluates the impacts of logging on NGOs, seeks to internalize global externalities biodiversity in tropical forests based on a into national environmental planning and the thorough review of the literature and on the Bank's sector work, operations, and its best expert opinion. dialogue with governments and partners. It is an iterative process, combining conceptual The study was commissioned from the Wildlife studies, reviews of state-of-the-art techniques Conservation Society (WCS). A draft paper was for measuring and mitigating local, national prepared by WCS and the University of Florida and global externalities, and testing these and was distributed to an expert review panel. concepts and tools in country-level studies as a Supported by the Forest Policy Implementation means of identifying good practices for Review and Strategy Process, these reviewers country planners and Bank task managers. The convened in Washington with the authors and results will help guide national actions to Bank staff in late February, 1999 for a conserve biodiversity, reduce greenhouse gas productive two day workshop after which the emissions, and protect international waters. paper was revised. The Overlays add a new dimension to Forests will continue to be harvested and timber traditional sector economic planning by harvesting will continue to be an important analyzing environmental impacts and component of sustainable forest management. opportunities to internalize global This is a reality we cannot escape. Therefore, as externalities. This analysis launches the this practice continues we need to assure "Impact Studies" which asks the broader ourselves that it is ecologically and question of what the positive and negative environmentally sustainable. The World Bank impacts of human activities are on has a responsibility to its clients to inform them biodiversity. An analytical framework is used of best practices and what the potential which allows for the systematization of these consequences of natural resources management impacts across sectors and areas of particular decisions are. This paper helps fulfill this interest. responsibility. Biodiversity Series - Impact Studies v Executive Summary Existing parks and protected areas are logging methods, collateral damage, and cornerstones of biodiversity conservation but on silvicultural approaches. Using the richness their own are inadequate to assure the present in both these terms, a framework for continued existence of a vast proportion of considering the impacts of logging and other tropical forest biodiversity. As a result, priority forest management activities on the various must be given to ensuring that the greatest components and attributes of biodiversity is possible amount of biodiversity is conserved presented. This framework is, in turn, used to outside protected areas by altering harvest evaluate the extensive literature covering patterns in these landscapes of resource different studies of logging in tropical forests. extraction. Of all forest uses within tropical forest regions, logging or timber harvesting, is findings: the most important to influence because not only is it the most lucrative, it also causes the 1. All consumptive uses affect some most severe direct and indirect environmental component or attribute of biodiversity, impacts. commonly affecting not only the target resource but other elements as well. Evaluating the impacts of logging on 2. As a result, only fully protected areas will biodiversity in tropical forests is not as simple a conserve all components and attributes of task as many would believe. It depends on a biodiversity. thorough review of the literature and on the 3. Recognizing that all significant best expert opinion due to the complexities interventions in natural forests have hidden under the seemingly simple rubrics biodiversity impacts, all silvicultural "logging" and "biodiversity." As a first step in decisions necessarily represent effectively analyzing the relationship between compromises. Management for some goods biodiversity and logging, it is essential to begin or services necessarily involves by disaggregating the terms "logging" and management against others. "biodiversity." 4. Different intensities and spatial patterns of timber harvesting, along with other silvicultural treatments, result in different effects on the different components of "biodiversity" into components (landscapes, biodiversity. ecosystems, communities, species/populations, 5. Some components and attributes of and genes) and attributes (structure, biodiversity are more sensitive than others composition, and function). It then to forest management activities. disaggregates "logging" by detailing the vast 6. There are some forested areas and some range of activities subsumed under this term areas within forests, which should never be including variation of logging intensities, logged. Biodiversity Series- Impact Studies vii Biodiversity Conservation in the Context of Tropical Forest Management 7. Biodiversity objectives can and should be areas of high priority for biodiversity established within production forests. protection, the persistence of poor logging 8. In many cases the logging itself is not the practices despite substantial efforts in research major cause of biodiversity loss, but rather and training, and the generally slow rate at the indirect effects, often promoted by the which most loggers are transforming presence of roads, (for example, hunting, themselves from timber exploiters into forest forest fires, and the likelihood of managers. deforestation) represent the major environmental threats. Nevertheless, even harshly treated forests maintain more biodiversity than oil palm Unfortunately, due to the complexities hidden plantations, cattle pastures, or cornfields. From under the seemingly simple rubrics "logging" a biodiversity maintenance perspective, natural and "biodiversity", the answer to the question forest management is preferable to virtually all "Is logging compatible with biodiversity land-use practices other than complete protection?" can only be the very unsatisfying protection. Forests that are carefully managed "It depends." The paper does not conclude for timber will not replace protected areas as with uncritical support for sustainable forest storehouses of biodiversity, but they can be an management of timber as a conservation integral component of a conservation strategy strategy. Such an endorsement is unwarranted that encompasses a larger portion of the given widespread illegal logging in the tropics, landscape than is likely to be set aside for strict widespread frontier logging and logging of protection. viii Environment Department Papers 1 Introduction Existing parks and protected areas are essential within managed forests should be protected, for biodiversity conservation but inadequate on and which approaches to silviculture to follow their own to assure the continued existence of within forests used for timber production the majority of natural landscapes, ecosystems, should be made on the basis of a wide range of communities, species and genotypes in tropical considerations. Forests differ in their forests. Even if the.oft-quoted goal of 10-12 biodiversity value, in their capacity to support percent protection were attained and the different intensities of silvicultural use, in reserved areas were appropriately located and pressures for conversion to non-forest use, and properly managed, up to 50 percent of tropical in the abilities of the relevant institutions to species would be expected to go extinct during regulate their management. General conceptual the next few decades (Soule and Sanjayan 1998). approaches to zoning forests for different uses Similar but often less well documented losses of have been presented recently by various authors the other components of biodiversity are including Noble and Dirzo (1997), and expected. Another way of considering this dire Frumhoff and Losos (1998). Methods for prediction is that if biodiversity outside of integrating conservation functions at different protected areas is neglected, thousands of geographical scales were reviewed recently by species are likely to disappear. Faced with this Poiani and others (2000). This paper focuses on bleak future, the first priority should be to the forests zoned for timber production. The increase the area of forests under strict goals are: protection while at the same time improving reserve management. Mechanisms should be 1. To contribute to the development of a developed to halt road building and commercial heuristic construct for considering the range logging in forest wilderness areas as well as in of impacts of forestry activities on tropical centers of diversity and endemism. However, forest biodiversity at. the levels of promoting more biodiversity-sensitive landscapes, ecosystems, communities, management of forests outside protected areas species, and genes. is of almost equal priority, given the 2. To indicate the topics on which further conservation potential of these still vast areas. research will be particularly useful in Biodiversity conservation within forests evaluating the impacts of different forestry managed primarily for timber production, for activities on the various components and example, should be fostered by a combination of attributes of biodiversity. not logging ecologically important areas and by 3. To suggest ways to mitigate the deleterious employing biodiversity-sensitive management impacts of forestry activities on tropical methods within production areas. forestbiodiversity. Decisions about which large forest areas should In the search for land-use practices compatible be designated for strict protection, which areas with biodiversity maintenance, many Biodiversity Series - Impact Studies Biodiversity Conservation in the Context of Tropical Forest Management environmentalists have focused upon logging motivate other conservationists to continue [see Grieser Johns (1997) and Haworth (1999) holding sustainable forest management (SFM) for comprehensive reviews of the environmental as a worthy conservation goal in forests outside impacts of logging in tropical forests]. This of protected areas (for example, Dickinson and emphasis is not surprising given that of all others 1996, Chazdon 1998, Poore and others forest uses, logging is often the most financially 1999, and Whitmore 1999). lucrative and has the most severe environmental impacts. The indirect effects of To help elucidate some of the factors upon logging, particularly increased hunting (for which the compatibility of tropical forest example, Robinson and others 1999) and the logging and biodiversity protection depend, this increased likelihood of deforestation due to paper first attempts to disaggregate the terms improved access, reviewed by Kaimowitz and "logging" and "biodiversity." The wide range of Angelsen (1998), have also been highlighted logging intensities, logging methods, collateral recently. What has emerged from the dust, mud, damage, and silvicultural approaches chainsaw noise, and diesel fumes is the idea appropriate for tropical forests is discussed. A that conservation of some components of framework for considering the impacts of biodiversity could be facilitated by logging and other forest management activities collaboration between loggers and on the various components (=landscapes, environmentalists. Some of the latter oppose the ecosystems, communities, species/populations, and idea of promoting better forest management as genes) and attributes (=structure, composition, and a means for achieving overall conservation function) of biodiversity is presented. In the goals (or at least oppose financial investment in main section of the paper, the components and such efforts, Rice and others 1997, Bawa and attributes of biodiversity are used to review the Seidler 1998, Bowles and others 1998). This impacts of logging and other silvicultural opposition notwithstanding, consideration of activities on tropical forests. To promote the inevitability of logging in much of the readability of the text, much of the tropics, the many constraints on expansion of bibliographical review is presented in nature reserves in tropical countries, the appendices organized by the components and challenges of protecting and managing the attributes of biodiversity. The paper concludes parks already demarcated on paper, sovereignty by suggesting ways to enhance the issues, development needs, and the implicit compatibility of forest management for timber adoption of the "use it or lose it" assumption, and biodiversity maintenance in tropical forests. 2 Environment Department Papers 2 Disaggregating "Biodiversity" Biodiversity refers to the natural variety and the physical organization or pattern of the variability among living organisms, the elements. Composition refers to the identity and ecological complexes in which they naturally variety of elements in each of the biodiversity occur, and the ways in which they interact with components. Function refers to ecological and each other and with the physical environment. evolutionary processes acting among the This definition and the elucidation below is elements. based upon OTA (1987), Noss (1990) and Redford and Richter (1999). Climate, geology Diversity of the landscape component refers to and physiography all exert considerable regional mosaics of land uses, land forms, and influence on broad spatial patterns of biotic ecosystem types. Structure of this component variety; local ecosystems and their biological could be described on the basis of the areas of components are further modified by different habitat patches, through the perimeter- environmental variation (such as local climatic area relations of these habitat patches, or on the and streamflow fluctuations) and ecological basis of interpatch linkages. Landscape interactions. This natural variety and variability composition pertains to the identity, distribution, is distinguished from biotic patterns or and proportions of different habitat types. conditions formed under the influence of Landscapefunction refers to issues such as patch human-mediated species introductions and persistence and interpatch flows of energy, substantially human-altered environmental species, and other resources. processes and selection regimes (Bailey 1996, Noss and Cooperrider 1994). Diversity of the ecosystem component refers to In discussions of diversity there is always the interactions between members of a biological lurking danger of assuming that "more is community and their abiotic environment. better." Were this the case, plowing roads Structure of this component might be measured through mature forests, clearcutting patches, through vegetative biomass and soil structural and introducing alien species would all be properties. Ecosystem composition refers to reasonable. Although decisions about which features such as biogeochemical standing species or community-types are most valuable stocks. Ecosystemjiunction pertains to processes are not straightforward, rarity is one obvious including biogeochemical and hydrological dimension that needs to be considered. cycling. Biological diversity can be measured in terms of Diversity of the community component refers to the different components (landscape, ecosystem, guilds, functional groups, and patch types community, population/species, and genetic), occurring in the same area and strongly each of which has structural, compositional, and interacting through trophic and spatial biotic functional attributes (Table 1). Structure refers to relationships. Structure of this component Biodiversity Series - Impact Sthdies 3 Biodiversity Conservation in the Context of Tropical Forest Management Table I Components and attributes of tropical forest biodiversity that might be influenced by logging and other silvicultural activities Organized by the components and attributes of biodiversity Components Struaure Composition Function Landscape Size and spatial distribution of Identity, distribution, and Habitat patch persistence and habitat patches (e.g., seral proportion of habitat types turnover rates; energy flow rates; stage diversity and area); and multi-habitat landscape disturbance processes (e.g., extent, physiognomy; perimeter-area types; collective patterns of frequency, and intensity of fires); relations; patch juxtaposition species distributions human land use trends; erosion and connectivity; fragmentation rates; geomorphic and hydrologic processes Ecosystem Soil (substrate) characteristics; Biogeochemical stocks; Biogeochemical and hydrological vegetation biomass, basal area lifeform proportions cycling; energy flux; productivity; and vertical complexity, density flows of species between patches; and distribution of snags and local climate impacts fallen logs Community Foliage density and layering; Relative abundance of Patch dynamics and other canopy openness and gap species and guilds; richness successional processes; proportions; trophic and food and diversity indices; colonization and extinction rates; web structures proportions of endemic, pollination, herbivory, parasitism, exotic, threatened, and seed dispersal and predation rates; endangered species; phenology proportions of specialists vs. generalists Speciesl Sex and age/size ratios; range Species abundance Demographic processes (e.g., Population and dispersion; infraspecific distributions, biomass, or survivorship, fertility, recruitment, morphological variation density; frequency; and dispersal); growth rates; importance or cover value phenology 6enetic Effective population size; Allelic diversity; presence of Gene flow; inbreeding depression; heterozygosity; polymorphisms; rare alleles; frequency of rates of outbreeding, genetic drift generation overlap; heritability deleterious alletes and mutation; selection intensity; dysgenic selection Note: Modified from Noss (1990) and Redford and Richter (1999). includes consideration of vegetative be measured through population age structure physiognomy and trophic structure. or species abundance distributions. Species/ Community composition refers to relative population composition pertains to which abundances of species and guilds. Community particular species are present. Function of this functions include flows between patch types, component refers to demographic processes disturbance regimes (such as fires and floods), such as recruitment and death. successional processes, and species interactions. Diversity of the genetic component refers to the Diversity of the species/population component refers variability within a species, as measured by the to the variety of living species and their variation in genes within a particular species, component populations at the local, regional, or subspecies, or population. Structure of this global scale. Structure of this component might component can be expressed on the basis of 4 Environment Department Papers Disaggregating "Biodiversity" heterozygosity or genetic distances between and in what proportions. Genetic functions can populations in different patches (that is, be expressed on the basis of gene flow, genetic metapopulation genetic structure). Genetic drift, or loss of allelic diversity in small, isolated composition refers to which alleles are present populations. Biodiversity Series - Impact Studies 5 3 Disaggregating "Logging" When properly planned and conducted, logging about the compatibility of logging with (=timber harvesting) is an integral component biodiversity conservation are complicated of forest management systems designed to because logging is carried out over a huge range promote sustained timber yields (STY), or the of intensities, using a variety of techniques more all-encompassing goal of sustainable which may be applied carefully or in ways that forest management (SFM). Unfortunately, result in a great deal of avoidable damage. The logging in tropical forests all-too-often following sections focus on the issues of represents a timber "mining" activity carried harvesting intensities, yarding methods (=how out without regard for the renewability of this timber is extracted from the stump to haul natural resource (for example, Putz and others roads), and ways of reducing logging damage. 2000). Other silvicultural treatments designed to A brief overview of other silvicultural promote sustainability are often prescribed but treatments used in timber stand management is are rarely applied. Due mostly to the desire to also provided. obtain voluntary third-party certification of good management from the Forest Stewardship Logging intensities Council (FSC), the transition from forest mining Trying to conclude anything about the to forest management has finally started to Try ity oflogging aboutethe occur in some areas in the tropics (Nittler and compatibility of logging and biodiversity Nash 1999). Despite this progress, even within maintenance is made complicated by the wide certified forests there are questions about how range of logging intensities, yarding methods, to most effectively and efficiently minimize the and accompanying forest management practices deleterious environmental impacts of logging to which tropical forests are subjected. The and other silvicultural activities. This paper simple observation that logging intensities span addresses many of these questions by reviewing more than two orders of magnitude (<1 m3/ha the state of knowledge about biodiversity to >100 m3/ha) illustrates the challenge of maintenance in exploited and managed tropical generalizing (Figure 1). forests. Even within the same forest management unit, It is critical to recognize that forest interventions logging intensities vary greatly at different of all types, from harvesting of fruits for home spatial scales. At a small scale (1-10 ha), the consumption to clearcutting for timber, all have typical aggregated distributions of tropical trees impacts on forests that need to be understood (Hubbell 1979) leads to locally severe logging and often deserve mitigation (Peters 1996). impacts unless harvesting controls are Nevertheless, logging is often the most implemented (that is, "tree-marking rules" are damaging and generally the most financially followed, Uhl and Vieira 1989, Crome and lucrative of such forest interventions (Pearce others 1992, Cannon and others 1994). The and others 1999). Unfortunately, discussions localized but severe direct impacts of roads, log Biodiversity Series - Impact Studies 7 Biodiversity Conservation in the Context of Tropical Forest Management landings, and skid trails hardly need to be eastern portions of Amazonian Brazil, it is likely emphasized (Guariguata and Dupuy 1997). At a that 30-50 m3 of 20-30 species would be slightly larger scale (10-100 ha), stands vary in harvested Uohns and others 1996). stocking of commercial species and in their accessibility due to terrain or edaphic factors. The substantial number of studies conducted on Where logging is carried out in areas designated the impacts of logging in tropical forests all for each year of management (=annual coupes), concluded that soil impacts and damage to the logging impacts tend to be aggregated at larger residual forest all increase with increasing scales. logging intensity (Ewel and Conde 1976, Sist and others 1998a). Proportions of both soil and Logging intensities also vary over time, tending residual trees damaged by logging range from 5 to increase with local timber shortages, to 50 percent, depending on harvesting improved access, and greater willingness of intensity, yarding method, and the care with markets to accept lesser-known species which the operations are carried out. (Plumptre 1996). For example, in a remote part Interpretation of data pertaining to the of the Bolivian Amazon loggers may harvest an relationship between logging intensity and average of only 0.3 m3/ha of 1-3 species residual stand damage is complicated by (Gullison and Hardner 1993) whereas in concomitant change in residual stand density; at similarly stocked stands in the more accessible the extreme, there is no residual stand in Figure I Logging intensities (m3/ha) for tropical forests. In most of these studies, as in most logging areas in the tropics, felling was with chainsaws and yarding with bulldozers or articulated skidders with rubber tires. < I rn/ha 50 m3/ha 00 m 3/ha 150 m 3/ha Bolivia Venezuela Indonesia Malaysia (Gullison and Hardner 1993) (Kammesheidt 1998) (Bertault and Sist 1997) (Pinard and Putz 1996) Suriname Costa Rica Philippines (Hendrison 1990) (Webb 1998) (Nicholson 1979) Venezuela (Mason 1996) Australia (Cromeet al. 1992) Brazil Malaysia Oohns et al. 1996) (Pinard and Putz 1996) Australia (Crome et al. 1997) Uganda (Chapman and Chapman 1997) 8 Environment Departnent Papers Disaggregating "Logging" clearcuts. Further complicating assumptions at the landscape, ecosystem, community, about logging damage is the fact that few population/species, and genetic levels varies tropical forests are logged (or burned) only once greatly along the continuum of technological (Box 1). sophistication of yarding methods that stretches from manual extraction to the use of helicopters Log yarding methods (Figure 2). Note that Figure 2 hides a great deal In this paper, particular attention is paid to the of relevant variation. In particular, attempts to yarding phase of the timber harvesting process include another dimension expressing yarding because much of the direct damage to tropical costs (per cubic meter yarded to the roadside) forest caused by logging occurs while logs are have so far failed due to the multitude of factors being extracted from the stump to roads or that need to be considered, including labor riversides from which they are then hauled or costs, technical capacity, and discount rates, towed out of the forest. Yarding methods along with the values assigned to future harvest utilized in tropical forests range in technological yields, non-timber forest products, biodiversity sophistication, the collateral damages with protection, and ecosystem services. which they are associated, and in yarding costs (Conway 1982). Accurate cost figures for Yarding can be carried out in an environmen- yarding are difficult to obtain and problematic tally/silviculturally sensitive manner, or can be to calculate (for example, should impacts on extremely destructive. The rankings of future timber yields be included, and, if so, at environmental impacts on Figure 2 are just what discount rate?). For the same volume of suggestions of the typical amounts of damage timber yarded, the range in biodiversity impacts associated with different yarding techniques. Box 1 Few tropical forests are logged or burned only once Logging and wildfires in tropical forests are causally linked (Holdsworth and Uhl 1997) and also share the characteristic of being self-promoting processes. Although the environmental damage resulting from unplanned logging of primary forest by untrained crews is justifiably receiving attention from researchers and policy- makers, few logged stands are allowed to recover for the full cutting cycle or rotation stipulated in their man- agement plans. Instead, forests are repeatedly entered by loggers who sequentially 'high-grade" the best re- maining timber. Although the costs of road building are substantial, the first cut is generally the most lucrative. As more timber species enter commercial markets, and mills begin to accept smaller and lower quality logs, incentives for multiple entry logging increase. The same logging firms are sometimes involved in a series of entries into the same forest, but more often firms with lower operating costs and smaller equipment take advantage of the increased access provided during the first harvest. A familiar adage in Amazonia is "ma- hogany builds the roads." Reliable data on frequencies of multiple entry logging, either legal due to waivers issued by governmental agencies or illegal, are apparently rare, but the process is unquestionably common- place. As with multiple entry logging, wildfires seldom occur only once (Peres 1999, Cochrane and Schulze 1999). In forests that are not naturally maintained by fire, the first fire opens the canopy and thereby subjects the under- story to rapid drying. Fuel mass in the understory is augmented by trees killed by the first fire, and by grasses and other plants that regenerate under conditions of reduced competition. The resulting fire frequencies far exceed the historical rates. In the eastern Amazon, for example, fire frequencies in Paragominas have recently doubled and now occur far too frequently to allow forest recovery. Human made savannas and grasslands along with severely degraded forests due to frequent fires are becoming more the rule than the exception throughout much of the wet and moist tropics. On the basis of either the area affected or the magnitude of impacts, the direct damages attributable to tropical forestry are minor compared to those of wildfires. Biodiversity Series - Impact Studies 9 Biodiversity Conservation in the Context of Tropical Forest Management Figure 2 Generalized ranges of direct biodiversity impacts of timber yarding methods used for tropical forest logging as a function of level of technological sophistication required (assumes equal volumes of timber yarded). For mechanized ground-based yarding operations, conventional (non-RIL) and reduced-impact logging (RIL) impacts are contrasted. Severe M odest Low (;reat Technological Sophistication For ground-based yarding, using the smallest bulldozer yarding, especially at high intensities yarding equipment possible contributes to on steep slopes logged during wet weather by reducing the deleterious impacts of logging, untrained crews is well known, but bulldozers W.Ahich yarding equipment is used is important, still yard much of the timber in commercial but so is the care with which yarding operations logging areas in the tropics. are carried out, as indicated by the contrasts between conventional and reduced-impact With the loss of forest in accessible areas in logging (RIL) on Figure 2. Unfortunately, much of the tropics, logging is increasingly although the silvicultural, environmental, and being relegated to flooded, steep, rocky, or economic benefits of planning of log extraction otherwise adverse terrain. Because ground routes have been recognized for many decades based yarding from such sites can be (see Putz and others 2000 for a review), well prohibitively expensive, they have often been planned logging operations are still the left as unlogged refugia within harvested areas. exception in tropical forests. Their status as refuges is jeopardized by helicopter yarding, because helicopters can yard Most destructive yarding in tropical forests is timber from even the most adverse sites. carried out with bulldozers (=crawler tractors). Obtaining timber from these sites is becoming Bulldozers are excellent devices for constructing more cost effective under some conditions. roads but are unfortunately versatile enough to Although areas from which timber is harvested yard timber as well. The excessive damage to by helicopters are not dissected by skid trails, soils and residual trees during conventional haul roads are still needed, and with them, all 10 Environment Department Papers Disaggregating "Logging" the problems associated with increased access. regeneration, the second cut releases the An equally important concern about helicopter established seedlings and saplings. There are yarding is that areas traditionally avoided by innumerable variations on the themes of loggers are rendered accessible and are likely to monocyclic and polycyclic harvesting, and each be harvested. Because helicopters leave no is appropriate under different conditions and in obvious trails, it is going to be very challenging forests for which there are different silvicultural to monitor their harvesting impacts. goals (Lamprecht 1989). Even within the same forest, the silviculturally most appropriate Logging and other silvicultural treatments harvesting regime sometimes can change over Logging can be either a cause of a great deal of distances of less than 100 m (Pinard and others avoidable damage or one of a series of 1999). silvicultural treatments designed to promote the regeneration and growth of commercial timber The most common method attempted for species while protecting ecosystem services and controlling timber harvesting in tropical forests biodiversity. In logging areas where sustained is to simply set a minimum stem diameter for yield of timber is a priority, various silvicultural felling. Although theoretically easy to treatments can be used in combination with the implement and monitor, minimum diameter appropriate logging regime to promote the rules are often inimical to achieving silvicultural regeneration or growth of commercial species. goals. The problem with this system is obvious Carrying out silvicultural treatments in where minimum diameter limits are set below conjunction with logging reduces their cost the sizes at which trees start to reproduce while simultaneously reinforcing the idea that (Appanah and Manof 1991, Plumptre 1995). logging itself can be silviculturally useful. Minimum diameter rules also do not prevent harvesting of clusters of trees and thereby Silviculturally appropriate harvesting creating silviculturally unsatisfactory conditions prescriptions (from a timber stand management such as where commercial species are favored perspective) run the full gamut from single tree by small canopy gaps or vines proliferate in selection to clearcutting (Smith and others large ones. 1997). For forests with ample stocks of trees of all sizes of the commercial species (that is, Logging is often the most severe of silvicultural negative exponential size-class frequency interventions, but there are other prescriptions distributions), "polycyclic" (=uneven aged) designed to increase the stocking of commercial methods such as single tree and group selection tree species, or to increase the growth of trees methods are generally suitable. In such a forest, already present. Retaining seed trees in the next crop is derived from the mixture of harvested stands is one possible way to increase trees of intermediate size (for example, 20-40 stocking, but for species that require mineral cm dbh) at the time of first cutting. In contrast, soil seed beds or minimal competition for at the time of the initial harvest the trees in the germination, establishment, and subsequent next crop in "monocyclic" (=even-aged) systems growth, seed tree retention needs to be are seeds, seedlings, or saplings. Clearcutting is combined with various other treatments. Seed the most obvious example of a monocyclic beds can be modified and competition can be system, but in "shelterwood management" a reduced by controlled burning, mechanical single-aged stand develops after two stages of scarification, or herbiciding plants competing harvesting. The first shelterwood harvest opens with seedlings of the crop species. Where the canopy and otherwise stimulates natural regeneration fails, or where particularly Biodiversity Series -Impact Stutdies 11 Biodiversity Conservation in the Context of Tropical Forest Management high stocking levels are desired, many foresters Reducing the impacts of logging and other have tried planting seedlings of commercial silvicultural treatments species in gaps or along lines cleared through Substantial attention has been given recently to the forest (see Box 2). For stands in which Suced-ial loggin has Box 3ie Most o regeneration of the commercial species is "reduced-impact logging" (RIL, Box 3). Most of already established, various thinning and weed the practices in the various RIL guidelines that control treatments are often prescribed to have been promulgated of late have long been accelerate tree growth and otherwise for "stand recognized as being environmentally sound and improvement." Thinning around potential crop silviculturally appropriate (Bryant 1914). Full trees, often referred to by tropical foresters as application of RIL techniques would represent a "liberation thinning," and vine cutting are two major step towards sustainable forest commonly prescribed but less commonly management (SFM), but RIL alone does not applied silvicultural treatments. In guarantee sustainability. Especially where tree experimental areas where liberation treatments species being harvested only regenerate in large have been applied to enhance volume clearings, the required silvicultural increments of commercial species, treatments interventions to assure sustained yield of the are often prescribed at 10-year or more frequent same species may often be substantial (for intervals (de Graaf and others 1999). As in the example, mimicking the impacts of slash-and- case of logging, all of these other silvicultural burn agriculture or hurricanes followed by fires, treatments have impacts on biodiversity, but Snook (1996), Fredericksen (1998), Dickinson they have been much less well studied. and Whigham (1999), Pinard and others (1999), Box 2 Enrichment planting Despite scarcity of successes and millions of dollars wasted, enrichment planting in gaps or along cleared lines continues to be invoked as a way of restoring the commercial timber production potential of severely degraded forests. The principal problem with enrichment planting is failure to tend the out-planted seedlings (Dawkins 1961, Dawkins & Philip 1998). Admittedly there are cases where few silvicultural options remain. Unfortu- nately, enrichment planting is often utilized in forests that should not need such an expensive intervention. Furthermore, it is often not recognized that enrichment planting as often applied results in conversion of natural forests into plantations. Mason and Thiollay (in press) compared the impacts of enrichment planting with conventional selective logging in Venezuela and found the former to have greater negative impacts on birds. With these caveats issued, a description of a large-scale enrichment planting project in Sabah, Malaysia may be useful because the project implementers are avoiding some of the pitfalls of this technique and are trying to mitigate some of its deleterious environmental impacts. Funds from the FACE (Forests Absorbing Carbon Dioxide Emissions) Foundation of The Netherlands, Rakyat Berjaya (a subsidiary of the Innoprise Foundation) have been used to plant nursery grown seedlings of com- mercial species in several thousand hectares of forests degraded by extremely intensive and poorly controlled logging in Sabah, Malaysia. The project has developed rapidly since its inception in 1992. Contract planters are now trained in dendrology so that they can recognize natural regeneration, and are paid for its retention at the same rate as for planted seedlings. Planters are also responsible for tending operations and are not paid for seedlings that do not survive 3 years. Site matching of species has also improved, as have nursery techniques for tending wildlings (=seedlings extracted from the forest) and raising large numbers of seedlings at rela- tively low cost. Finally, to promote retention of wildlife in the planting area, seedlings of fleshy- fruited trees are planted along with the mostly wind dispersed timber species in the Dipterocarpaceae. When the planted stands are ready to harvest in 50 or so years, the concessionaire hopes to harvest 50-100 trees per hectare, an intensity tantamount to clearcutting but perhaps preferable to the forest conversion to pulpwood plantation option that has been adopted in much of the region. 12 Environment Department Papers Disaggregating "Logging" Box3 Is reduced-impact logging (RIL) cheaper? Many people believe that RIL has been proven cheaper than conventional logging (CL) and are therefore per- plexed at the lack of its adoption by loggers. Adoption of RIL techniques is of particular importance in the tropics where few countries have regulations that are both explicit enough and well enough enforced to pre- vent exploitative harvesting. Numerous studies over the past decades have shown that by planning skid trails and directional felling to facilitate yarding, logging damage can be substantially reduced (Bryant 1914, Nicholson 1958, Redhead 1960, Fox 1968, Nicholson 1979, Ewel and Conde 1980, Hendrison 1990, Johns and others 1996, Pinard and Putz 1996, Blate 1997, Elias 1997, Winkler 1997, Uhl and others 1997, Haworth 1999). Several experimental plot-based studies have demonstrated that due primarily to reduced yarding costs, such straightforward improvements in logging methods also translate into direct financial benefits to the loggers (Marn and Jonkers 1981, de Graaf 1986, Malvas 1987, Jonkers 1987, Hendrison 1990, Gerwing and others 1996, Bertault and Sist 1997, Barreto and others 1998, Holmes and others 1999, Boltz 1999). If RIL is cheaper than CL per cubic meter yarded to the roadside, then why haven't loggers spontaneously adopted RIL techniques out of enlightened self interest? A partial answer to this question may be that under normal operating conditions, at industrial scales, and espe- cially on adverse terrain in wet forests, RIL may not always be cheaper (Putz and others 2000, Hammond and others 2000). Data on the comparative costs of RIL and CL from the "Reduced-Impact Logging as a Carbon Offset Project" in Sabah, Malaysia may serve to elucidate this issue. Tay (1999) and Healey and others (in press) reported that the financial profits from logging were substantially lower in a 450 hectare area harvested ac- cording to RIL guidelines than in a comparable area subjected to CL. The principal reason for lower profitabil- ity of RIL was the reduction in yield caused by reductions in the proportion of the area logged due to restric- tions on bulldozer access to steep slopes. Despite regulations against timber yarding from steep slopes in Sabah, the practice is commonplace and loggers consider the timber foregone when RIL guidelines are fol- lowed as lost profits and thus do not spontaneously adopt RIL out of enlightened self interest. These findings indicate that the issue of cost effectiveness of RIL under adverse conditions deserves scrutiny and that loggers not using RIL techniques may not be acting in a financially irrational manner. In contrast to the apparent situation regarding RIL adoption in Southeast Asia, large vertically integrated forestry companies in Brazil have begun to invest heavily in RIL apparently out of enlightened self-interest. Although there is an additional incentive for adoption of more environmentally friendly logging in the form of better enforcement of regulations governing forest management operations, it appears that some companies have been convinced by the work of Baretto and others (1998), Holmes and others (1999), and others that their I profits will increase if they adopt more efficient (and resource friendly) practices. Loggers in the region are also applying RIL techniques in response to an increase in demand for timber certified by the Forest Stewardship Council. Given the differences in the results of comparative studies on the financial profitability of different logging techniques, adoption of RIL techniques under some conditions may require either subsidies (such as carbon offset funds) or stricter regulations with better enforcement. The conditions under which incentives for RIL adoption are needed, and the best form and magnitude of these incentives are yet to be determined. Fredericksen and Mostacedo (2000). This reduced as well, by proper design, construction, silvicultural challenge notwithstanding, careful and maintenance of road networks. Much of the planning and implementation of harvesting cost of harvesting timber and a large proportion guidelines would represent a big step towards of the hydrological damage (for example, sustainable forest management. stream sedimentation) due to logging is associated with roads (Bruijnzeel 1992). For For all yarding methods, logging damage can be ground-based yarding, efficient skid trail substantially reduced, and logging costs layouts are also essential for reducing logging Biodiversity Series -Impact Studies 13 Biodiversity Conservation in the Context of Tropical Forest Management damage and increasing yarding efficiency, but environmental impacts, adhering to RIL improper road siting often restricts skid trails to guidelines has advantages regardless of inappropriate places. Although the required whether the forest is allowed to regenerate or is density of roads is lower where aerial extraction replaced by an oil palm plantation or a maize techniques are used (such as skyline and field (Congdon and Herbohn 1993, Nussbaum helicopter yarding), roads are still needed and and others 1995, Pinard and others 1996). their locations can greatly influence logging costs and environmental damage. Guidelines The impacts of other silvicultural treatments on for road construction are abundantly available biodiversity (such as thinning and vine-cutting) and are well outlined in the FAO Model Code of depend on the intensity with which they are Forest Harvesting Practices (Dykstra and applied and on the proper designation of areas Heinrich 1996). Unfortunately, forest deemed inappropriate for stand "improve- engineering standards in most tropical logging ment." For example, vinecutting can enhance areas are extremely low and there are too few tree growth but undoubtedly has negative experienced forest engineers involved in most impacts on a wide variety of animals (Putz and tropical logging operations. It is important to others in press). Both biodiversity impacts and note that because soil damage due to poor skid labor costs of vine cutting depend on whether trail design or improper use, for example, only selected future crop trees are liberated or reduces productivity, increases surface erosion, vine cutting is carried out as a blanket and has various other deleterious prescription. 14 Environment Department Papers Imnpacts of Forest Management on Biodiversity Background exploitation (Coomes 1995, Homma 1992). This ahistorical and wishful thinking is extremely Human activities are highly variable in their angerocau a llows its adhrentlt influence on the components and attributes of dneosbcuei losisahrnst binfleer ony theompone ant thattribults of believe that there exist easy, cost-free solutions biodiversity. Any human activity that results intmxliaino h lnt substantial resource extraction or modification t e will always entail significant, often unknown, As societal concerns about the fates of tropical and almost always unappreciated consequences forests increase and human demands on forests for one or more biodiversity components, primarily by redirecting matter and energy change, so should the ways in which they are flows. The cumulative redirection is enormous silviculturally treated. For example, a few decades back when the primary component of at the planetary scale as three examples sustainability of interest to most forest decision- illustrate: 1) Vitousek and others (1997) calculated that 40 percent of the Earth's makers was sustained timber yields (STY), it terrestrial primnary productivity is being was often recommended that logging be appropriated by humans, 2) 25 to 35 percent of followed by silvicultural stand "improvement" the primary productivity of continental shelf treatments such as poison girdling of non- marine ecosystems is consumed by humans commercial trees, for a historical review see (Roberts 1997), and 3) Postel and others (1996) Dawkins and Philip (1998). Unfortunately, few report that humans now appropriate 26 percent of the tens of thousands of hectares of tropical of total evapotranspiration and use 54 percent forests that were treated according to the of all runoff in rivers, lakes, and other accessible various silvicultural systems used in the 1950s sources of water. (for example, the Malayan Uniform System and the Tropical Shelterwood System) remain, most Despite these statistics on current human having already been logged again, converted to impacts and the observations of the long-term oil palm plantations, or otherwise lost. impacts of previous generations of our species Fortunately, there is now evidence from a (Denevan 1992), many in the conservation and lowland dipterocarp forest in Malaysia that at sustainable development community still least the Malayan Uniform System (MUS) could maintain it is possible to both use and preserve achieve its silvicultural goals (Lee and others biodiversity (Huston 1993) with no costs to 1998) with mixed impacts on biodiversity, either side. This claim is made regardless of a depending on the taxon in question. The forest history of human over-exploitation of resources treated by MUS and then studied 45 years later that began in prehistory (Goudie 1990) and is was, as intended, enriched in commercial manifested most recently in the negative effects timber trees, but also maintained much of the of tropical logging (Bawa and Seidler 1998, fauna and flora of primary forest. More studies Frumhoff 1995) and non-timber forest product of this sort are needed, and they need to be Biodiversity Series - Impact Studies 15 Biodiversity Conservation in the Context of Tropical Forest Management better circulated so that more people will realize hydrologic processes. The most severe impacts that at least "qualified" successes in tropical described in this section, however, result from silviculture are attainable. indirect consequences of logging such as increased access to remote areas, fragmentation, In the following sections (4.2-4.6) the impacts of and altered fire regimes. logging and other silvicultural treatments on the components and attributes of biodiversity using Structure: The size and spatial distribution of the framework elaborated in section 2.0 (see tropical forest patches and the juxtaposition and Table 1) are outlined; details are provided in the connectivity of different forest patches across appendices. Summarizing the impacts of the necape are most patche across forestry activities on biodiversity in tropical the landscape are most affected by ite indirect forests is an effort fraught with problems in impacts of logging. One of these impacts- large part due to their immense diversity. Even increased access to humans-is often at the species level, the diversity is difficult to deleterious. Partcularly when logging roads imagine and each of the literally millions of penetrate far into the forest frontier in countries species in tropical forests responds to different where there are numerous potential forest logging impacts in distinct ways. While some colonists, logging is almost invariably general response patterns are obvious, and accompanied by increased hunting pressure others have emerged from field research, the (Robinson and others 1999, Robinson and idiosyncrasies and apparent inconsistencies of Bennett 2000) and is often followed by species responses need to be recognized. For deforestation as lands are cleared for agriculture example, chimpanzee (Pan troglodytes) (Kaimowitz and Angelsen 1998). populations have been reported to increase (Howard 1991, Hashimoto 1995), decrease Another indirect impact of logging on this (White 1992), and not respond to logging attribute is fragmentation of previously (Plumptre and Reynolds 1994, see Appendix contiguous or otherwise connected forest VI). In contrast, studies of terrestrial and bark- patches. The resulting forest fragments, gleaning insectivorous birds have consistently however, are typicaly not completely isolated reported negative impacts of logging (see for all species for all time. For example, wide Appendix IV). Many more examples of this sort lor roads may time. Forossable are discussed briefly below, and both cosset logging roads may represent uncrossable aredisconsistent pattefly below,erge. consistent barriers for some forest interior species but and inconsistent patterns emerge. roadsides with secondary vegetation attract Landscape impacts (see Appendix I) many large ungulates where they are more easily hunted (Robinson and Bennett (2000) in Logging affects the landscape component ofSaak,Mlyi)Whtelogdsnsar biodiersit by cangig lan form andSarawak, Malaysia). Whether loggred stands are biodivrsity y chaging lnd fors andinterspersed within s ecies-rich forest or in a ecosystem types across large geographic areas p p (Box 4). Logging shifts the regional mosaic of low diversity landscape dominated, for land uses. Although the landscape component example, by pulpwood plantations, also of biodiversity is the least sensitive to logging, influences long-term species maintenance. changes in the size, spatial distribution, and Furthermore, even small unlogged patches connectivity of habitat patches across the within harvest areas can serve as source landscape occur especially as the intensity of populations for some species after logging. The management interventions increases. These degree of fragmentation depends on whether changes in the habitat mosaic alter species logging is dispersed over large areas, or is distribution patterns, forest turnover rates, and concentrated in small areas (Figure 3). Where 16 Environment Department Papers Impacts of Forest Management on Biodiversity Box4 The critical role of production forests in maintaining biodiversity in the tropics Tropical rainforests harbor approximately 50 percent of all terrestrial biodiversity and much of that biodiversity occurs in the lowland forests which are most accessible for, and thereby most threatened by, logging and agricultural conversion. In Peninsular Malaysia, for example more than 50 percent of all mammal species occur below 350 m and a startling 80 percent occur below 650 m (Stevens 1968, MacKinnon and others 1996). World- wide tropical forest loss is estimated as at least 17 million hectares/yr, an area the size of Cambodia (FAO 1993). Dramatic as it is, this global total is probably a gross underestimate given that recent figures show that Indonesia alone has lost 18 million hectares of tropical forests between 1985 and 1997 in just three of the major Outer Islands-Kalimantan, Sumatra and Sulawesi. Most of this forest loss has occurred in lowland forest and logging has played a major role in that destruction. In many areas forests have been cleared and converted for agriculture, plantations and transmigration. However, it is not logging activities per se that have caused forest and biodiversity loss but poor concession management and new access along logging roads. These factors have allowed shifting farmers and hunters to colonize and clear logged areas. Moreover, areas that have been logged (whether legally or illegally) are more vulnerable to wild fires during El Nifio years; there is no doubt that poor logging practices contributed to the great Indonesian fires in 1998 when 5 million hectares of forest in Kalimantan burned, an ecological, economic and social disaster. It is not just the reduction in total area of natural habitat that concerns conservationists, it is also the fact that remaining habitat is being broken into ever smaller fragments within which species' populations are no longer viable. As habitats become more fragmented by clearance along new roads, protected areas will increasingly become "islands" and more and more species populations will be fated for local extinction. Fragmentation is particularly serious because few protected areas still cover a full range of altitudinal habitats as large areas of lowland forests, the most species-rich habitats and prime habitat for many large and wide-ranging mammals, have been excised from designated protected areas for logging (such as Kerinci, and Gunung Leuser National Parks in Sumatra). For Gunong Leuser National Park, for example, the main elephant, orangutan and tiger populations occur in the lower-lying production forests outside present park boundaries. Even more alarming is the recent increase in logging and other illegal activities within the boundaries of several National parks in Indonesia. The loss of bird species on Java is an indicator of what can be expected in Borneo and other large islands as forests are cut and fragmented. Studies of forest bird distribution shows that whereas in Borneo, Sumatra, and New Guinea, maximum avian species richness is still found in lowland forests and species numbers decrease with altitude, Java is atypical in having a depauperate avian fauna in the hill zone between 300 and 15OOrn. This pattern is interpreted as a result of long term deforestation since some predominantly lowland species have failed to survive in hill forests because they have been cut off from lower altitude populations which were a source of colonizers. Families of large birds such as malkohas have lost proportionally more species than families of small birds such as flowerpeckers. Moreover, Java has lost more species which are exclusive to lowland rainforests than species which can occupy secondary habitats, forest edge or open habitats. Small forest patches have lost more species than larger blocks, with extinction rates as high as 80 percent in 10-40 hectare plots compared to rates of 25 percent for areas over 10,000 hectares. These results have important implications for reserve design, forest management and biodiversity conservation. They emphasize thatforests must be maintained asforests and that reserves must be large, cover a wide altitudinal range and be connected by corridors of natural habitats. This necessitates a landscape approach to biodiversity conservation by manag- ing production forests adjacent to core protected areas to maintain both permanent forest cover and biodiversity so that production forests become buffer zones that effectively extend and supplement the conservation estate (MacKinnon and others 1996, MacKinnon and Phillipps 1993). preservation of forest interior biodiversity is the conservation biologists (Noble and Dirzo 1997). priority, concentrating logging in small areas is Fortunately, due to associated cost savings, generally the pattern recommended by concentration of logging activities is one of the Biodiversity Series - Impact Studies 17 Biodiversity Conservation in the Context of Tropical Forest Management Figure 3 Some impacts of logging on biodiversity as a function of distribution of logging activities (percent of area logged) and logging intensity (m3/ha of timber harvested) Concentrated Damage Worst Case for Biodiversity .t .to Moderate Yield Maximum Yield Large Area Impact-Free Small Area Impact-Free Do C Low Impact Dispersed Damage J ? Minimum Yield Moderate Yield Large Area Impact-Free Small Area Impact-Free Low High Area Logged (percentage) steps towards improved forest management improved access provided by logging roads that loggers may find acceptable. indirectly fosters post-logging habitat changes by human forest colonizers, weeds, and The impacts of logging and other timber stand wildfires. management activities on landscape structure are minimized when plantation products are Setting aside reserves within logging areas may used in place of those from natural forest. mitigate some of the deleterious impacts of Opting for this so-called "New Zealand logging and other silvicultural treatments. The Solution" (Hunter 1998) is not currently an specific location of reserves substantially option likely to be exercised in many tropical influences their value in biodiversity countries. Nevertheless, as forests dwindle and conservation. Optimally, the full range of economies grow (such as in Thailand, Malaysia, landscape features and habitats should be Chile, and South Africa), this solution will represented within protected areas. become more viable. Function: Logging may markedly alter several Composition: Logging activities may directly landscape level ecological processes subsumed and indirectly affect the identity, distribution, under the functional attribute of the landscape and proportion of habitat types in tropical component of biodiversity. For example, forests. Forestry may directly affect composition logging roads, and activities associated with of the landscape component of biodiversity by their construction, can greatly influence the intentional creation of new types of habitats permanence of the forest fragments they create (such as forests converted into plantations). by altering landscape level disturbance regimes. Furthermore, if silvicultural objectives are In large part, disturbance is altered because uniform across the landscape, inter-stand roads and skid trails provide ready access to the diversity is sacrificed by widespread forest for both colonists and fire (see below). application of the same stand "improvement" High intensity and widespread logging, treatments. Perhaps most importantly, especially if not carefully controlled, also 18 Environment Department Papers Impacts of Forest Management on Biodiversity influences hydrological processes at all levels improvements in relevant remote-sensing perhaps including the regional climate. Where methods, the impacts of the understory burns logging roads are wide and logging intensities typical for tropical forest fires are substantial. are high, landscape level movements of animals Especially in forests that have not burned for can be disrupted (Goosem 1997), as can gene many decades to centuries, the effects of even a flow in plants when pollinators are restricted to single understory burn are great, even if they isolated fragments by inhospitable are not fully realized for many years. Typically surroundings. It should be noted, however, that small trees are killed whereas large trees often the impacts of roads depends on where and receive only minor basal damage. Unfortu- how they are constructed, and change with time nately, basal damage exposes trees to wood as the road and its margins mature (Lugo and rotting organisms; a high incidence of heart rot Gucinski 2000). is typical of forests that burn at low intensities at infrequent intervals (F. E. Putz, pers. obs.). In this section the emphasis is on the impact of Elevated rates of tree mortality rates for many fires on the functional attributes of landscapes years post-fire are also characteristic in seldom- in full recognition of the importance of fires to burned forests. Such long-term data are difficult all components and attributes of tropical forest to obtain because accumulation of fuel in the biodiversity. Although fires have long played form of fallen branches as well as post-fire substantial roles in the ecology of tropical proliferation of palms, grasses, and other fire- forests throughout the world (Goldammer 1990, carrying plants render forests very fire prone. Saldarriaga and West 1986), wildfires as Surprisingly, the effects of tropical forest fires on exemplified by the recent Indonesian fires have biodiversity have not been extensively studied recently increased in frequency, extent, and (Leighton and Wirawan, 1986). intensity in part due to widespread logging Ecosystem impacts (see Appendix II) (Cochrane and Schulze 1999, Cochrane and others 1999). Canopy opening due to timber The deleterious ecosystem-level impacts of harvesting operations increases the rate of forest logging on tropical forests are widespread, drying and thereby increases inflammability substantial, enduring, and well studied even in characteristically wet forests and (especially compared to the landscape and swamps (Holdsworth and Uhl 1997). Perhaps genetic components). The ecosystem component equally importantly, vegetation proliferation of biodiversity is somewhat more sensitive to along logging roads and skid trails greatly logging impacts than the landscape component enhances fire penetration into forest interiors in part because management activities are even where logging damage is slight or non- usually implemented at this scale. In contrast to existent. Whether or not the extensive forest the landscape component, most ecosystem-level fires that have recently become common can be impacts are a direct consequence of logging indirectly attributed to logging, it is clear that activities. Logging purposely removes biomass fire damage often exceeds logging damage in from ecosystems, but it also alters their vertical many tropical countries. In Bolivia in 1999, for complexity and soil characteristics. Depending example, fires burned an estimated 20,000 km2 on the silvicultural objectives, changes in whereas logging was carried out in perhaps structural heterogeneity may be intended. only one-tenth of that area (W. Cordero, pers. Whether intended or not, the structural impacts comm.). of logging alter the relative proportions of lifeforms and biogeochemical stocks, as well as Although difficult to detect on satellite images, nutrient and hydrologic cycling, productivity but see Souza and Barreto (in press) for recent and energy flows. Biodiversity Series - Impact Studies 19 Biodiversity Conservation in the Context of Tropical Forest Management Structure: Logging may affect the structural lowest logging intensities, to substantially attribute of the ecosystem component of greater amounts where heavy logging is biodiversity by changing the biophysical followed by poison girdling of non-commercial properties of soils, spatial heterogeneity of trees in the residual stand. Necromass, forest stands, and biomass. The extent and types including coarse woody debris, increases of ecosystem damage to these structural immediately after logging but may then attributes depend on logging intensity, yarding decrease to levels below pre-logging conditions system, and the care with which the operations due to increased temperatures near the ground are conducted. Soil compaction, for example, is and associated increases in decomposition rates. a major problem during ground-based yarding Wildfires promoted by logging road operations especially where skid trails are construction and canopy opening, as well as unplanned and yarding continues during wet controlled bums carried out for silvicultural weather. Where compaction is severe, soil purposes, can have obvious effects on biomass permeability and bulk density often require and necromass stocks in tropical forests. many decades to recover. Exposure of mineral soil after litter layers and root mats are bladed Maintenance of healthy communities, species off by bulldozers is also a concern. Mineral soil populations, and gene pools is predicated upon disruption during bridge building, road protection of hydrological functions, nutrient construction and maintenance, and skidding cycles, and other ecosystem properties. operations also represent forest management Fortunately, methods for mitigating the activities that affect ecosystem structure and ecosystem-level impacts of logging on tropical thereby have biodiversity impacts. forests are well known. Switching from ground- based to skyline yarding techniques, for Logging induced soil compaction negatively example, greatly reduces damage to residual affects hydrology, which in turn alters the stands, soils, and streams but allows harvest on characteristics of watercourses. For example, steep slopes. The opposite trend in tech- water infiltration rates into soil, surface flow nological change also can have environmental rates and volumes, and stream channel benefits; log yarding with draft animals or by characteristics are all adversely affected by manual means generally results in substantially uncontrolled logging. Deposition of large less logging damage than yarding with volumes of unconsolidated sediments into bulldozers (Cordero 1995). To enjoy these streams during road construction and due to potential benefits, the expertise of experienced increased erosion from exposed mineral soils on forest engineers should be more often called log extraction routes can also greatly modify upon where logging does have to occur. stream characteristics such as pool-riffle-run ratios. The impacts of such changes on aquatic Composition: One important way that logging organisms have been little studied in the tropics. affects the ecosystem-level compositional attribute of biodiversity is by changing Another impact of logging on ecosystem-level biogeochemical stocks. For example, soil structure is reduction in biomass and alteration compaction reduces water holding capacity, of necromass. Losses of biomass due to forest which in turn leads to increased surface runoff. management activities include the amounts in Limited storage capacity in natural streams is the removed timber, damage to trees in the further reduced by sedimentation, which means residual stand that result in mortality, and any that flow regimes can be greatly modified by silvicultural treatments that result in tree death. logging, especially during the first years after Biomass losses range from 5-10 Mg/ha at the logging is completed. Various RIL techniques, 20 Environment Department Papers Impacts of Forest Management on Biodiversity such as installation of cross drains on skid trails, release. The deleterious impacts of logging on can greatly diminish these impacts. forest carbon balance can be greatly diminished by application of RIL techniques (Pinard and Where large volumes of timber are harvested Putz 1996). Substantial biodiversity benefits are and post-logging forest recovery is retarded by also likely to result from RIL, but this and other soil damage, fire, or weed infestations, standing co-benefits have not been well studied. Other stocks of nutrients in biomass are greatly silvicultural treatments such as fire reduced. Storage of nutrients released from management, weed control, thinning, and biomass in necromass and soil organic matter is enrichment planting have various impacts on generally brief in lowland tropical forests due to both greenhouse gas emissions and biodiversity accelerated rates of decomposition and low that should be investigated. cation-exchange capacities in the soil. The results can be substantial leaching and, if the forest is burned, volatilization of nitrogen and Logging, especially if followed by silvicultural sulfur. treatments such as liberation of future crop trees from competition, can substantially change the Function: Logging affects the functional physiognomy, composition, and trophic attribute of ecosystem-level biodiversity by structure of forest stands. To a large extent, adversely affecting hydrological and these modifications represent the goal of forest biogeochemical fluxes as well as productivity. "refinement" treatments applied to increase Reduced plant productivity results in part from volume increments and relative densities of impeded root growth, a consequence of logging- commercial timber species. This "stand induced soil compaction. Because most of the domestication" by nature reduces species available nutrients are usually found near the richness; rare, threatened and endangered top of the soil profile, blading of the soil surface species may become locally extinct especially if also diminishes nutrient availability in local they have no perceived commercial value. areas and otherwise interferes with nutrient These changes in composition and structure cycling. In more extensive portions of logging affect numerous community-level ecological areas where RIL guidelines are not followed, processes including colonization, predation and nutrient cycling and hydrological functions are mortality rates, pollination, seed dispersal, and greatly modified by reduced canopy timing and abundance of flower and fruit interception of rain and mist, decreased uptake production. of water and nutrients by the diminished Structure: The most obvious logging-induced biomass, and increased occurrence of surface impact on the structural attribute of erosion and landslides especially associated community-level biodiversity is the change in with improperly located and poorly constructed proportions of successional stages in forest roads and skid trails. stands. Depending on harvesting intensities, planning of roads and skid trails, and training Changes in carbon storage and flux associated and supervision of workers, logging can result directly and indirectly with logging and other in large changes in the proportion of forest in silvicultural activities influence whether forests mature, recovering, and early successional are net sources or sinks of "greenhouse" gases. stages. In some severely disturbed areas, For example, substantial logging-induced succession might be "arrested" by post-logging transfers of living trees to coarse woody debris proliferation of vines, bamboo, and other non- can have substantial effects on understory arboreal growth forms. Silvicultural treatments structure and dynamics leading to more carbon such as thinning and vine-cutting can increase Biodiversity Series - Impact Studies 21 Biodiversity Conservation in the Context of Tropical Forest Management the rate of succession and increase the microclimates that result from exploitation, proportion of stand growth concentrated in silvicultural treatment, and by hunting (see Box 5). commercial species, but not without affecting Species impacts (see Appendices V and VI) biodiversity in more than the intended ways. The species component of biodiversity has Composition: For the community component of received the most attention from researchers biodiversity, logging affects composition by concerned about the impacts of logging and changing (often purposefully) the relative other silvicultural treatments in tropical forests. abundance of species and guilds inhabiting The most obvious species-level impact of forest stands. Relative abundances of tree logging is on the abundance and age/size species with light demanding vs. shade tolerant distribution of harvested and damaged trees. regeneration, wind vs. animal dispersed seeds, Depending on the intensity of logging and the vertebrate vs. invertebrate pollinated flowers, care with which it is carried out, the and thick vs. thin bark, for example, are all reproduction, growth, and survival of a great subject to change in logged and otherwise number of species can be adversely affected. In silviculturally treated forests. Likewise, reviewing this literature, it is important to note sieviculturatiyntreatedfforest. uikewofanise, s that the taxa studied were not selected at rersntto odifrnguilsonml random. Instead, in many cases, the species (such as understory insectivores and arboreal chosenswer d, to be csensitie to,cand folivores) is influenced by forestry activities,. thus goo d i a of fensitr impacs Depending on a great number of factors related to the intensities, spatial scales, and modes of Structure: The most immediate and direct forest intervention as well as characteristics of impacts of logging on the structural attribute of the focal taxa, effects of forestry activities can be the species component of biodiversity are negative, positive, or neutral. For example, in suffered by the harvested tree species. Their eight studies that considered the impacts of populations are often left greatly depleted, logging on frugivorous birds, two reported especially in the larger size classes of positive impacts at the guild level, three reproductive individuals when management is reported negative impacts, and three reported based solely on minimum diameter felling no change at all (Appendix IV). rules. Because of the spatial clustering characteristic of many commercial timber trees, Function: The functional attribute of the richest patches of forest are generally the community-level biodiversity includes most severely disturbed, unless logging numerous key ecological processes (such as guidelines specify minimum spacing between pollination, herbivory, seed dispersal and harvested trees. predation) all of which are affected by logging Changes in forest structure are suffered most by especially under the most intensive specialist species of the forest interior. After management interventions. Many of the effects logging, many formerly shaded microenviron- on these processes are a direct consequence of ments in the forest interior become drier, altered resource abundance (for example fruit brighter, warmer, and more easily exploited by for frugivores or young leaves for folivores), some predators. For example, severe canopy which in turn result from the logging-induced opening adversely affects litter invertebrates changes in community structure and and their predators. For species that are composition. In addition to being influenced by generalists in their diets and wide-ranging in resource-base changes, these ecological their habitat use, such as many frugivorous processes are also affected by changes in forest canopy birds, the direct impacts of logging vary 22 Environment Department Papers Impacts of Forest Management on Biodiversity Box 5 Hunting in logged tropical forests The escalating scale of wildlife harvest in logged areas is an insidious problem that may undermine the biodiversity conservation potential of forests managed for wood production (Robinson and others 1999, Fimbel and others in press). Logging operations multiply the harvest of wildlife from tropical forests primarily by increasing access to previously remote areas. These same access roads serve as conduits for commercially traded meat and other wildlife products. Although the magnitude of the impacts of subsistence hunting of tropical forest animals (for food, feathers, and skins for example) by indigenous people are non-trivial (Robinson and Redford 1991, Redford 1992, Robinson and Bodmer 1999, Peres 2000), the impacts of commercial hunting are several orders of magnitude greater. Of particular concern is the tendency for hunters to target almost any species larger than 1 kg, including many species that are vulnerable to extinction due to their long life spans and low rates of population recovery (Bodmer and others 1997). Hunting pressures increases in logging areas partially because of the number of people involved in logging operations as well as by the increased access provided by logging roads. In formerly isolated forest areas, loggers themselves hunt (Ruiz and others in press) or provide ready markets for meat and other wildlife products (Wilkie and others 1992, Bennett and Gumal in press). Local communities quickly shift toward com- mercial hunting to supply markets to which logging roads provide access. This shift initiates a positive feed- back cycle wherein money gained from wildlife products buys better weapons with which to hunt, which in tum increases the harvest. Such escalations are unsustainable, however, and some species become locally ex- tinct. Loss of wildlife threatens the sustainability of tropical to the extent that the animals targeted by hunters play key roles in ecological processes including seed dispersal, seed predation, and herbivory (Redford 1992, Jansen and Zuidema in press). Even when animal species are not extirpated, their numbers may be reduced suffi- ciently such that they are rendered ecologically extinct. The loss of wildlife may have cascading effects on the structure and composition of the tropical tree community with deleterious consequences for recruitment of commercial timber species. Such dire predictions are most likely to be borne out in areas such as the Guyanas where a large proportion of canopy trees are dispersed by large animals (Hammond and others 1996). Finally, as human populations expand and forest landscapes become more fragmented, the potential for wildlife to re- colonize defaunated areas will likely diminish. While the deleterious consequences for wildlife of logging and other silvicultural treatments (such as vine cutting) deserve further investigation and mitigation, it should be recognized that the indirect impacts of log- ging resulting from increased access often far outweigh the initial damage done by even the worst predatory logging. from being somewhat negative, to neutral, to Species impacts of logging and stand refinement positive. For the understory species that are treatments are of particular concern in small adversely affected by logging, the effects may forest management units. Private landowners persist for decades (Wong 1985, but see Lee and with less than 100 hectares to manage, for others 1998). It is worth reiterating that the example, may be unwilling to set aside 10 imnpacts of hunting on populations of large and percent of their forest for species preservation. If slow-reproducing animals generally overwhelm their forests are surrounded by similarly managed or deforested areas, then blanket any potential benefits that they might have apation o refiement treatmet application of stand refinement treatments or enjoyed after the canopy was opened by logging heavy logging can take a heavy toll on (Bennet and Dahaban 1995, see Box 5). commercial and non-commercial species alike. Population sizes and structures of most species are also drastically modified by the fires that so Composition: Logging affects the composition often accompany uncontrolled logging. attribute of species-level biodiversity by Biodiversity Series - Impact Studies 23 Biodiversity Conservation in the Context of Tropical Forest Management changing the abundance and distribution of across or even within taxa. Our review of the species. Unless logging is accompanied by other literature on primates, for example, revealed silvicultural treatments designed to foster their few cases of consistent responses of species to reproduction and growth, the abundance and logging (Appendix VI). This variation can be population structure of the harvested tree attributed to differences in logging intensities as species are greatly modified by logging (see Box well as to differences in the duration of post- 6). Logging impacts on tree populations logging population monitoring. Silvicultural continue for many years after logging is treatments other than logging, especially vine completed because damaged trees suffer high cutting and crown liberation of future crop mortality rates, proliferation of weeds (such as trees, might have more consistent deleterious vines) interferes with tree reproduction and impacts on canopy animals, but such impacts survival, and population size reduction and have been little studied. fragmentation can decrease pollination levels and change patterns and intensities of seed dispersal Small fragments of untouched forest that and predation. Species composition of animals also remain within even heavily logged forests serve changes in response to the direct impacts of logging as important refugia for plants and animals. (such as canopy opening) and the associated Wildlife densities in these unlogged fragments indirect impacts as well (for example increased fire can be very high during and shortly after frequency and intensity, hunting, and forest harvesting, but then diminish as animals conversion). recolonize the surrounding matrix. Many 1munplanned" reserves are on steep or otherwise Changes in species composition in response to adverse sites, which certainly influences their forestry operations are by no means consistent function as refugia. Implementation of Box 6 Lesser-known species-Marketing challenges, silvicultural impediments, or desirable diversity? Sustaining volumetric yields of timber is made challenging in tropical forests where many trees have little market value (Plumptre 1996). Particularly where just a few species of light-demanding trees have ready markets, species that are commercially lesser known or that have less desirable timber properties often render regeneration-enhancing treatments prohibitively expensive (Fredericksen 1998, Pinard and others 1999). Where markets for formerly lesser-known species have developed and they can therefore be harvested profitably, forest managers can financially justify creating the large canopy openings required for regeneration of the most valuable light-demanding timber species. The same lesser known species that interfere with timber stand management treatments and present chal- lenges for wood technologists and marketing firms represent an important component of forest diversity. One indication of their importance is that whereas tree species with wind dispersed seeds (such as Dipterocarpaceae and Meliaceae) dominate the international timber trade (ITTO 1996), a high proportion of species producing less well known timbers produce fleshy fruit and animal dispersed seeds (Jansen and Zuidema in press). Silvicultural treatments applied to promote regeneration and growth of commercially desirable species might be made more feasible by increased market demand for what are currently lesser known timbers. While pro- moting sustained yield of the species being harvested, these silvicultural treatments intentionally result in at least locally substantial modification of pre-intervention stand structure and composition. Compromises be- tween sustained yield and the more all-encompassing goal of sustainable forest management need to be in- formed by research on the direct and indirect impacts of increased timber harvesting (Fredericksen and others 1999). 24 Environment Department Papers Impacts of Forest Management on Biodiversity helicopter yarding, and to a lesser extent other management units in which population sizes of aerial yarding techniques, may place many of all species are correspondingly small. Allelic these unlogged patches in jeopardy. frequencies of commercial species change after removal of a large proportion of healthy Function: Demographic processes (such as reproductive adults. For species with high survivorship, fertility, and recruitment) and densities of advanced regeneration, genetic growth rates are two key functional attributes of structure of their populations are unlikely to the species component of biodiversity that change dramatically after selective harvesting, logging affects. Populations of many organisms unless collateral damage is severe. Timber stand are susceptible to large fluctuations after improvement treatments also may affect the logging due to both the direct impacts on forest genetic structure of species targeted for removal conditions (for example microclimate and (such as woody vines) as wetr as their fragmentation) and the indirect impacts of (uha od ie)a ela hi fragmentingfio and forest condirectsim s Te associates, but these impacts have apparently hunting, fire, and forest conversion. The not been studied. Given the high proportion of proliferation of disturbance-adapted taxa in vines and other plants that resprout after logged-over forests, some species of which are cutting (=coppice), large impacts on genetic not native or were not previously common in cure are lkely. the area, can have large but as yet little studied structure are unlikely imnpacts on the resident flora and fauna. Composition: The fact that most species are rare Genetic impacts (see Appendix VII) in tropical forests implies that allelic diversity will decrease with increasingly intensive The genetic component of biodiversity is likely management interventions. Unregulated to be the most sensitive of all components to harvesting of all merchantable individuals of logging because of reductions in effective commercial species, for example, has immediate population size and interruptions in gene flow. impacts on allelic frequencies that continue to At present, however, little is known about the change due to decreased effective population genetic structure of any tropical organisms, sizes. Deleterious recessive genes may become even commercially valuable timber trees (Ledig more apparent due to dysgenic selection and 1992). Furthermore, the techniques required for heterozygosity may decline due to the assessing the genetic structure of populations "bottyeneck" effect in the small, isolated are sophisticated and expensive. Except in a few potions that inut smarvested cases, concerns about dysgenic selection, genetic populations that result from harvesting, forest drift, and other genetic problems are based on fragmentation, and other direct and indirect controversial theory that is rapidly developing impacts of forestry activities (Styles and Khosla as evidence accumulates. 1976, Murawski and others 1994a and b, but see Newton and others 1996). Structure: Logging affects the structural attribute of the genetic component of Function: Logging may affect the functional biodiversity by reducing effective population attribute of the genetic component of sizes and heterozygosity. Effective population biodiversity by interrupting gene flow, which in sizes of both commercial and non-commercial turn influences outbreeding rates. Decreased species are reduced by harvesting, other effective population sizes coupled with losses of silvicultural treatments, forest fragmentation, pollinators and seed dispersal agents can result weed proliferation, and wildfires. There are also in reduced gene flow and inbreeding depression good reasons to be concerned about the effects in populations of both commercial and non- of logging and stand improvement treatments commercial species. Especially vulnerable are on dioecious species and in small forest populations represented by scattered mature Biodiversity Series - Impact Studies 25 Biodiversity Conservation in the Context of Tropical Forest Management individuals and very few juveniles (for example obligate outcrossers, only very severe many "long-lived pioneers" such as the reductions in effective population size are likely mahoganies). Given the high proportion of to have much effect on gene flow (Ghazoul and tropical tree species that are dioecious or others 1998). 26 Environment Department Papers Overview of Biodiversity Con- servation in Relation to Logging 5 and other Silvicultural Treatments Synthesis The impact of these silvicultural approaches on A way of graphically displaying the impacts of each biodiversity component using three logging on tropical for-ests (Figure 4) based categories were assessed. First, each upon the various components and attributes of biodiversity component was scored as 'mostly biodiversity as described above was developed. conserved" for cases in which their attributes Along the vertical axis of our framework the are expected to usually stay within their natural five components of biodiversity were arrayed in range of variation. Second, biodiversity the order of increasing susceptibility to logging components were scored as "affected" for cases impacts (landscape, ecosystem, community, in which their attributes are expected to population/species, and genetic). The frequently fall outside their natural range of horizontal axis arrays a variety of approaches to variation. Finally, biodiversity components were silviculture in order of increasing intensity. scored as "mostly lost" for cases in which their Figure 4 Expected effects of a range of forest uses on the components of biodiversity ___- - .Legend: NTFP = Non-timber forest prod- . ~~~~~~~~~~~~~~~ucts Genetic _ fA > MQSTLYRIL = Reduced-impact logging c ~~~~~MOSTILY 3} .1% Reserves = Protected areas 5 Species -oa LOST within logged units 0 Refinement = Silvicultural E treatments such as liberation of future crop trees from competi- 6 Community AFFECTED tion, which can substantially change the physiognomy, com- ._% .position, and trophic structure L \ . of forest stands which are ap- > \csse plied to increase volume incre- .2 Ecosystem \ ' ments and relative densities of n0 MOSTLY \ commercial timber species 0 m CONSERVED Enrichment planting = Increas- Landscape ing the stocking of commercial species by planting seedlings (or seeds) in logging gaps or along A ~ ~ Yx x x cleared lines t~ - 900 CL = Conventional logging Biodiversity Series - Impact Studies 27 Biodiversity Conservation in the Context of Tropical Forest Management attributes are expected to almost always fall use capability, biodiversity value, or the outside their natural range of variation. capabilities and desires of local stakeholders. At least two other dimensions of this multi- The scoring process used for Figure 4 was dimensional topic are captured in Figure 5, on admittedly subjective; the scores were based on which it is indicated that every forest activity a reading of the literature and the authors' can be carried out over a range of intensities. experience. It is fully recognized that particular Correspondingly, each forest-use activity situations might warrant different scores, and it generates timber volumes and financial profits is emphasized that the purpose is to illustrate that also range widely (Figure 6). Recognizing what is believed to be a useful analytical that forest use practices vary over time with, for process rather than to obtain perfectly accurate example, market fluctuations and political scores. It is also emphasized that this change, and that biodiversity impacts are a framework serves to suggest hypotheses that function of a multitude of interacting factors seem to be important research topics for operating at different temporal and spatial conservation biologists from a number of scales, it is hoped that Figures 4-6 present an different disciplines. accurate "snapshot" of the relationship between biodiversity conservation and forest The responses summarized on Figure 4 actually management. represent a multitude of impacts from a diversity of silvicultural approaches applied The impacts summarized in Figure 4 and the with varying degrees of concern for biodiversity ranks and ranges of impacts and profits on over a large range of scales. Additionally, Figure Figure 5 are based on a combination of literature 4 addresses neither the issue of profitability of reviews and the authors' subjective estimations. different forest uses nor issues related to land- For example, under the range of stocking levels, Figure 5 Generalized biodiversity impacts plotted against expected short-term financial returns to forest owners or concessionaires (NTFPs = non-timber forest products) Severe (A et nv/e tona v oggin Y 1g (CL)/ E u Reced / 'hi~~~~~~~~~~~~~~~i .t E .0 Modest P areas Small Great Financial Profitability 28 Environment Department Papers Overview of Biodiversity Conservation in Relation to Logging and other Silvicultural Treatments Figure 6 Generalized biodiversity impacts resulting from different approaches to forest management for timber. (CL = conventional logging, RIL = reduced-impact logging, refinement = any of a variety of silvicultural treatments applied to increase rates of timber volume increment; and, reserves = protected areas within logged units) Severe E mn 4-. sia 1 Modest 0 1 2 3 4 5 6 Commercial Timber Production (m3/ha/yr) terrain, and accessibility that logging is carried 1) Tropical forests are often assumed to have out, reduced-impact and conventional logging developed their current structure, overlap in both impacts and profitability. composition, and functional properties Nevertheless, particularly where logging is under a disturbance regime characterized carried out on steep slopes or under otherwise by small, localized, and natural adverse conditions, application of most RIL perturbations. The importance of guidelines results in immediate profits that are unrecorded widespread cataclysmic lower than those enjoyed by unconstrained anthropogenic and natural disturbances, conventional loggers. This observation helps especially those of past centuries, is seldom explain why where regulations are non-existent recognized. or unenforced, conventional logging will likely be used to log such areas. Correspondingly, 2) It is often assumed that regenerating trees some of the impacts of high intensity RIL harvested for timber is simply a matter of overlap those of low intensity conventional protecting advanced regeneration (for logging, but the former generally has fewer example, seedlings, saplings, and poles) and adverse environmental impacts than the latter. providing small canopy openings in which Troubling assumptions they can mature. Three widely held and interlocking assumptions 3) Many people believe that devolution of about tropical forests complicate the issue of the authority over forests to indigenous groups compatibility of forest management for timber and other rural communities will enhance and biodiversity conservation: efforts towards sustainable management Biodiversity Series - Impact Stuidies 29 Biodiversity Conservation in the Context of Tropical Forest Management and biodiversity conservation (Luckert are less secure. Some communities, such as the 1999, Putz 2000). municipalities in Bolivia that have recently received control over substantial forest areas, While these assumptions hold true in some may be indifferent about conservation tropical forests, many of the trees currently (Kaimowitz and others 1998). In other cases, being harvested actually regenerated under such as some communities in Nepal, devolution conditions very different than those of today or of forest control has had substantial those that will be created by harvesting alone. conservation benefits. Obviously the process of The most familiar example is provided by the devolution deserves scrutiny lest some turn out neotropical and African mahoganies (Meliaceae to be worse forest stewards than the logging spp); these trees and many others like them are companies and national governmental agencies the heritage of slash-and-burn agriculture, they replace. Empirical evidence suggests that hurricanes, river meanders and/or catastrophic community behavior towards forests depends fires of centuries past. The problem confronting on the forest richness or forest poorness of their silviculturalists managing for such light- particular situation, but many other factors are demanding species is that promoting likely involved and deserve investigation. regeneration requires intensive stand manipulations which are expensive and have Conclusions great impacts on the biodiversity present All consumptive use affects some component or immediately prior to intervention (Lugo 1999). attribute of biodiversity, commonly affecting not only the target resource but other factors as well Despite accumulating evidence of substantial (Redford and Richter 1999). The population/ and long-term pre-historical and more recent species component is most commonly impacts of humans on what were formerly understood to be affected by silvicultural considered "natural" ecosystems, it is important activities although effects, some of which are to recognize that the current rates, spatial scales, subtle or cumulative, are undoubtedly often and intensities of human-induced perturbations missed even in this comparatively well studied often far exceed the resilience of tropical forests. component. Of increasing importance is an Forest recovery after disturbances such as understanding of how the community and mechanical clearing for pastures, for example, is ecosystem components have been (Runnels known to be extremely slow, especially if the 1995) and are being affected by logging and pastures are large or intensively used. Natural other silvicultural activities (Noss and regeneration in such areas can be accelerated Cooperrider 1994, Vitousek and others 1997, through application of silvicultural treatments, Fimbel and others, in press). but full recovery of diversity, from the genetic to the landscape level, will require centuries, if it Recognizing that all significant interventions in occurs at all. For example, although leaf natural forests have biodiversity impacts, all biomass may recover quickly, in abandoned silvicultural decisions necessarily represent agricultural clearings coarse woody debris compromises. Management for some goods or accumulates at rates measured in centuries. services necessarily involves management Research progress on methods for afforestation against some others. What are "weeds" to and reforestation should not beguile us into timber stand managers are food sources, rare believing that, once destroyed, tropical forests species, carbon stores, or inter-crown pathways can be recreated. for other human and non-human stakeholders. The biodiversity compromises involved in The cause of social equity may be well served deciding whether to cut vines, retain seed trees, by devolution of management responsibility to or enhance seedling establishment by carrying local communities, but the biodiversity benefits out controlled burns should be informed by 30 Environment Department Papers Overview of Biodiversity Conservation in Relation to Logging and other Silvicultural Treatments research. Unfortunately, very little is known and training, and the general slow rate at which about how tropical forests can be managed in most loggers are transforming themselves from the most biodiversity friendly manner. timber exploiters into forest managers. Researchers have instead focussed on Nevertheless, even the most harshly treated enumerating the deleterious environmental forests maintain more biodiversity than tree impacts of uncontrolled logging by untrained farms for pulpwood, oil palm plantations, and unsupervised crews. To inform decisions maize fields, or cattle pastures. Furthermore, about tropical forest management and to assure logging is often the environmentally least that biodiversity is protected to the maximum damaging of land uses that are also financially extent, more research is needed on how to viable (Pearce and others 1999). Given these maintain diversity in forests selected for conclusions, effective mechanisms for financing logging. The large and rapidly growing body of forest protection and environmentally sound literature on ecosystem management in both forest management are needed. It might also north and south temperate forests (such as prove useful to evaluate critically the limits to Kohm and Franklin 1997, Lindenmayer 1999) the assumption that well managed forests are should provide inspiration and starting points less likely to be converted to other land uses for tropical researchers intent on solving than forests that are logged without apparent biodiversity related problems associated with concern for sustainability. forest management. By focusing on the deleterious environmental The primary conclusions to derive from these impacts of tropical forest management analyses are that: activities, we often lose sight of the fact that from a biodiversity maintenance perspective, D Different intensities and spatial patterns of natural forest management (that is, maintaining timber harvesting, along with other forests as forests) is preferable to virtually all silvicultural treatments, result in different land-use practices other than complete effects on the different components of protection. The number of taxa with reportedly biodiversity. inconsistent, neutral, or positive responses to a Some components and attributes of logging is an indication that timber harvesting biodiversity are more sensitive than others is not necessarily incompatible with protection to forest management activities. of many components and attributes of * Only extremely limited use will protect all biodiversity. This conclusion derives even components (that is, large protected areas greater support from the fact that nearly all of are essential for biodiversity conservation). the studies conducted to date on the biodiversity impacts of logging were carried out The capacity to mitigate the deleterious after unplanned logging by crews with no environmental impacts of logging and other training in reduced-impact logging techniques silvicultural treatments should not be construed and no incentives to reduce their impacts. As as constituting unilateral support for forest management practices improve under sustainable forest management as a market pressure or pressure from land-owners, conservation strategy. Such an endorsement is the deleterious environmental impacts of unwarranted given widespread illegal logging logging and other silvicultural activities are in the tropics, widespread frontier logging and likely to be substantially reduced. Forests that logging of areas of high priority for biodiversity are carefully managed for timber will not protection, the persistence of poor logging replace protected areas as storehouses of practices despite substantial efforts in research biodiversity, but they can be an integral Biodiversity Series - Impact Studies 31 Biodiversity Conservation in the Context of Tropical Forest Management component of a conservation strategy that and effectively extend the conservation estate. encompasses a larger portion of the landscape Finally, it should be recognized that landscape than is likely to be set aside for strict protection. management is consistent with the ecosystem In other words, forests managed primarily for approach emphasized by the Convention on timber, if managed properly, will supplement Biological Diversity (CBD). 32 Environment Department Papers 6 Recommendations 1. Given that all substantial forest components of reduced-impact logging and interventions affect some component or various other pre- and post-logging attribute of biodiversity, the best way to silvicultural activities. Particular attention assure biodiversity conservation is through should be paid to plant and animal species the establishment and protection of large, with attributes that render them susceptible properly located, and well managed to logging-induced extirpation (Martini and reserves and protected areas. others 1994, Pinard and others 1999). Regional guidelines for certification need to 2. In some parts of the world and some types be developed and regularly updated to of forest, there should be no logging at all. reflect increasing knowledge about biodiversity so as to assure that certification 3. Existing legislation and regulations should efforts realize their conservation potential. be enforced. Where they do not exist, or are weak they should be created or updated to 7. Measures to protect biodiversity should be reflect current conservation and sustainable included in all forest management plans. resource management concerns. Particular attention should be paid to the 4. Within forests used for logging and other retention of nest and den trees as well as silvicultural activities, biodiversity species of great importance to biodiversity conservation is enhanced by setting aside a conservation. portion of the area for complete protection. 8. Wildfires need to be controlled. Successful Optimally, these reserves should be large, fir e contro lled. Sude shaped so as to minimize edge effects, cover fire control programs will need to include representative areas of all the ecosystem public awareness building as well as types present, and include features of implementation of existing technologies and special concern for biodiversity methods. Research is needed for developing maintenance such as water courses, rock cost-effective fire control measures with outcrops, and salt licks. minimal biodiversity impacts. 5. Where logging is to be carried out, reduced- 9. While continuing to promote and develop impact logging guidelines (Dykstra and community-based wildlife management Heinrich 1996) should be fully implemented prograrns, hunting by loggers and market by well trained crews. hunting of slow reproducing species should be prohibited. Enforcing existing laws will 6. Researchers need to determine the financial in many cases confer substantial protection and environmental costs and benefits of the to species threatened by over-hunting. Biodiversity Series - Impact Studies 33 Biodiversity Conservation in the Context of Tropical Forest Management 10. Training is needed for researchers who will 11. The biodiversity impacts of devolution, develop biodiversity-sensitive silvicultural plantation conversion, certification, methods as well as cost-effective methods 12. forest-based carbon offsets, and global for monitoring the environmental impacts trends in timber markets need to be of silviculture. Training is also needed for monitored. the field crews that are responsible for implementing the recommended practices. 13. Linkages to the policy arena must be Perhaps the most critical shortage is of pursued. Ecosystems provide important forestry/conservation "research services which typically are not internalized practitioners" who understand both the into the decision making process. broad scientific and the more specific Legislation and their regulations are an technical aspects of tropical forest integral part of this process and should be management. reviewed, updated and enforced. 34 Environment Department Papers Appendix I - Impacts of Logging (and other silviculture treatments where noted) on the Landscape Component of Biodiversity in Tropical Forests Biodiversity Series - Impact Studies 35 Impacts of logging (and other silviculture treatments where noted) on the landscape component of biodiversity in tropical forests LandscaPe attributes Category Structure Composition Function Vegetation, Soil & - Fragmentation / roads + greater access to > 50% forest area logged + species lost > Increased human colonization and fire & Water humans 4 increased deforestation (Kaimowitz & (Eastern Amazon: Nepstad et al. 1992) decreased migration of animals and gene flow Argelsen 1998) in plants (if pollinators are restricted to fragments) > Tree populations, biomass, and community > Ecological degradation of 1 °forest fragments > More than 50 yrs required for recovery of pre- composition affected more by edge effects from increased edge effects & new habitat logging structure (Uganda: Plumptre 1996) than by patch size (Amazonia: Gascon & matrix (Amazonia: Gascon & Lovejoy 1998) 200 yrs necessary for recovery of spp Lovejoy 1998) composi- tion following logging damage (Malaysia, forest gap model: Kurpick et al. 1997) Increased spatial heterogeneity of habitats after > Increase in mixed forest (w/ valuable timber o Impoundments due to undersized or failed 140 yrs of agroforestry, logging, & charcoal spp) & decrease of Cynometra forest patches culverts & bridge abutments kill trees and may production followed by 50 yrs regeneration after arboricide treatment from 1951-90 influence fauna (F.E. Putz, pers. obs.) (Puerto Rico: Garcia-Montiel & Scatena 1994) (Uganda: Plumptre 1996) )> Increased likelihood of large scale accidental > Maintenance & recovery of species fires due to canopy opening and roads composition in logged patches depends on (Amazonia: Cochrane & Schulze 1999) or fuel landscape context (Liu & Ashton 1999) accumulation from fire protection (Miombo woodlands, Africa: Chidumayo 1988) ;O Complex mosaic of forest types & disturbance > Harvest gaps warmer than natural treefall gaps w/ 85% swamp forest & only 15% lowland (Amazonia: Vitt et al. 1998) forest escaping logging disturbance (W. Kalimantan, 8 yrs post-logging: Cannon et al. 1994) > Patches (I - 100 ha) of unlogged forests in heavily logged area (500- 1000 ha, 80-120 m3/ha) on steep slopes (>350) and in rocky areas (Sabah, Malaysia: Pinard & Putz 1996) Primates Non-rodent mrommals Rodents & > Increased small mammal richness in forest Marsupials fragments b/c of increased edge and new habitat matrix Gascon & Lovejoy 1998) Birds Herps > Neotropical frogs - see rodents above (Gascon & Lovejoy 1998) Insects > Loss of species & genes of butterfly communities b/c of conversion of 30% of landscape to agriculture and forestry (Neotropics: Brown 1997) Soil organisms Appendix II impacts of Logging (and other silviculture treatments where noted) on the Ecosystem Component of Biodiversity in Tropical Forests Biodiversity Series - Impact Studies 39 Impacts of logging (and other silviculture treatments where noted) on the ecosystem component of biodiversity in tropical forests Ecosystem Attributes Category Structure Composition Function Vegetation, Soil & P Reduced biomass, basal area, and stem density > Thin barked species most susceptible to P Increased vulnerability to fire: ignition after 2 Water (Uhl et al. 1991, Silva et al. 1995, Pinard & Putz logging (and fire) (Amazonia: Uhl & Kaufman weeks w/out rain in gaps vs. months in 1996, Webb 1998, Finegan & Camacho 1999) 1990; Pinard & Huffman 1997) understory (Amazonia: Cochrane & Schulze > Reduction in large trees with hollows & holes 1999, Nepstad et al. 1999) important for wildlife (Australia: Gibbons & Lindenmayer 1996) > 2/3 basal area, 3/4 number undamaged trees, > Higher richness (trees > 20 cm dbh) in logged P Decreased evapotranspiration & infiltration; I/5 no of emergents (>30 m), & 1/3 vs. unlogged, but when equal numbers of trees increased surface flow 4 nutrient losses commercial volume of unlogged forest sampled richness was equal (W. Kalimantan, 8 (Brtower 1996, Poels 1987, Bruijnzeel 1992), but (Venezuela, 19 yrs post-harvest of 10 trees/ha: yrs post-logging: Cannon et al. 1998) nutrient bsses don't restrict productivity (Proctor Kammesheidt 1998) > T related to 1992) > Forest surface fires kill 40-95% of all trees w/ geography than whether a patch was logged; > Soil bulk density 6 yrs post-logging 60% greater dbh > 10 cm (Amazon: Nepstad pers obs) richness higher in west of reserve and in logged than in undisturbed forest (Malaysia: Malmer & patches (L.tanda, 60 yrs post 'sustainable logging': Grip 1990); skid trails estimated to require > 50 P 36% biomass loss from Increased treePkp-e16)yst co'rhdaiccduty(aly: motlt (Aaoi, IAA fro frgmn Plumptre 1996) yrs to recover hydraulic corductvity (MaVsia: mortality (Amazonia, I00 m from fragment ~&Knrzna 96 edge, 10-17 yrs post-fragmentation: Laurance K& K 1996) etal. 1997) > Increase in organic fuels (56 to 180 tons/ha) on > Reduced densities of timber spp & shift in > 100- 170 M tons of carbon/yr emitted from forest floor (eastern Amazon, post-logging: Uhl dominance & age structure of canopy spp 1981-1996 (Asia: Houghton & Hackler 1999) &Kaufman 1990, Uhl & Vieira 1989) (Puerto Rico, post 140 yrs agroforestry, > 0.5% of total global carbon emissions (Brazilian > Soil disturbance main factor limiting seedling logging, charcoal production & St yrs Amazon, 1996-97: Nepstad et al. 1999) regeneration & establishment (Kalimantan: regeneration: Garcia-Montiel & Scatena 1994) Gardingen et al. 1998) > 5-40% of forest floor disturbed by roads, skid > Epiphytic & strangler Ficus spp suffer high rates > Post CL, stream suspended solids and turbidity trails & tractor tracks (Cannon et al. 1994 of damage (Malaysia, post-logging: Lambert were 12x & 9x > control and levels persisted Johns et al. 1996, Pinard & Putz 1996, Sist et al. 1991) at least 5 yrs; RIL levels were 2x > control but 1998a, Holmes et al. 1999) recovered after 2 yrs (Malaysia: Zulkifli & Anhar 1994) Vegetation, Soil & > Soil properties (i.e. BD, total porosity, sat hydr > Woody vines increased w/in 100 m of edges > Watershed sediment loads were 1 8x higher 5 Water conductivity, & resistance to penetration) but didn't compensate for lost biomass from mos post & 3.6x higher I yr post-logging Vegetation required 12-52 yrs (skidtrails longest) to increased tree mortality Amazonia forest (Malaysia: Douglas et al. 1992) recover (Malaysia: Kamaruzaman & fragments 10-17 yrs pcst fragmentation: Laurance > 200 yrs necessary for recovery of spp comp Kamaruzaman 1996) et al. 1997) following logging damages (Malaysia, forest gap > Limiting harvest gaps to < 650 m2 (< 2 model: Kurpick et al. 1997) trees/gap) enhanced dipterocarp regen & > Runoff as % precipitation ranged from 58% - inhibited pioneer regen (Kalimantan: Gardingen 94% in plantations converted from forest et al. 1998) depending on method (Malaysia: Malmer 1992) > Harvest gaps much larger than natural treefall > Road & skid trail impoundments may increase gaps & thus have less litter and shade & more methane emissions (F.E. Putz, pers. obs.) intense light (Amazonia: Vitt et al. 1998) > Sediment loads have negative effects on > Tree basal area and commercial volumes dwsra qai ie(utai:Cmbl recovered within 40 yrs post logging w/ MUS downstream aquatic life (Australia: Campbell & (Malaysia: Manokaran 1998) Doeg 1989) Primates > Long distance seed dispersal and species movements to/from different habitat patches likely impeded (no specific data) Non-rodent > Species movements to/from different habitat mammals patches likely impeded (no specific data) Rodents & > Species movements to/from different habitat Marsupials patches likely impeded; seed predation rates likely altered (no specific data) Bats > Species use of different habitat patches & pollination & seed dispersal rates likely altered (no specific data) Birds > Bird spp decline w/ increasing disturbance > Species use of different habitat patches likely (Lawton et al. 1998) altered; depending on spp., pollination, predation, seed dispersal rates likely altered; (no specific data) Herps Community Attributes Category Structure Composition Function Insects > Arthropod diversity decreases when structural > Various ecosystem processes will be relatively complexity decreases (Holloway et al. 1992; unaffected (Vitousek 1990) Gardner et al. 1995) and/or b/c of changes in understory, litter, & soil microclimate (Blau 1980, Kremen 1992, Spitzer et al. 1997, Camilo & Zou in press) Soil organisms ) 25%-75% reduction in soil mycorrhizae > Termite richness lower in clear-cuts, but (Malaysia, post-logging: Alexander et al. 1992) higher in regenerating plots than I' forest (Cameroon), but overall richness was similar in 1 °, 3 yr and 17 yr secondary sites (Malaysia: Eggleton et al. 1995); changes in termite comp may be delayed after disturbance (Eggleton et al. 1997) > Nematodes declined along disturbance gradient & were 40% lower in clearcut vs. 1' forest (Bloemers et al. 1997) > Soil fungi increased from 17 spp in 7-yr regrowth to 27 in 16-yr regrowth; soil microbial popin's positively correlated w/ tree density & basal area (India: Arunachalam et al. 1997) > Ectomycorrhizal diversity greater under open & regenerating canopies than in undisturbed forest (Kalimantan, 9 months post clear-cut or partial logging, trees > 50 cm dbh: Ingleby et al. 1998) > Termite populations increase after conventional logging more than after RIL (Sabah, Malaysia, 80-120 m3/ha harvested: F.E. Putz, pers. obs.) (MUS = Malayan Uniform System, a monocyclic approach to timber stand management) Appendix III Impacts of Logging (and other silviculture treatments where noted) on the Community Component of Biodiversity in Tropical Forests Biodiversity Series - Impact Studies 43 Impacts of logging (and other silviculture treatments where noted) on the community component of biodiversity in tropical forests Community Attributes Category Structure Composition Function Vegetation > Increased proportion of open > Increased species richness (esp. light-demanding pioneer trees, shrubs i Decreased seed dispersal of 6 timber canopy (Uhl &Vieira 1989, and lianas) > 10 yrs post-logging (Brazil: Verissimo et al. 1995, species in logged + hunted forest vs. Thiollay 1992, Cannon et al. Magnusson et al. 1999; Salick et al. 1995, Silva et al. 1995, Delgado et al. logged + protected forest (Costa Rica: 1994, Mason 1996); up to 1997, Costa Rica: Webb 1998) Guariguata et al. in press) 50% in gaps vs. 5-10% D Aggressive shade-intolerant spp dominate and competitively exclude > Fruit production of trees lower IO yrs (Hartshor 1978, Denslow other plants (Uganda: Struhsaker 1997) post-logging (Amazonia: Johns 1992) > Greater density of > Multiple tree felling gaps most diverse, but dominated by pioneer tree > High degree of outcrossing and low density understory vegetation 5-6 spp (Costa Rica: Webb 1998) of mature individuals (neotropics: yrs post harvest (Venezuela: Murawski 1995, Stacy et al. 1996) Mason 1996) Remaining large trees o High dominance of few spp in heavily disturbed habitats (e.g., roads) > Seed production lower in several timber inadequate to sustain future (Costa Rica: Guariguata & Dupuy 1997) species (Uganda: Plumptre 1995) harvests (Philippines, I 5 yrs Large-scale, long-term shift in spp composition after logging, augmented - Increased self-pollination attributed to low post-SLS: Tombec & by arboricide treatment of non-commercial spp (Uganda: Plumptre density of flowering individuals (Murawski Mendoza 1998) 1996) & Hamrick 1991, 1992) 4 less seed > No change in overall richness & composition in logged vs. logged + thinned (Nicaragua; Salick et al. 1995) Reduced tree density & > Increased irradiance 4 more early successional spp (Swaine 1996, > Wind-dispersed seeds fall in higher canopy closure, denser Mulkey et al. 1996, Whitmore 1998) especially vines & pioneers (Lowe numbers in gaps than in understory cover < 2 m above ground, & Walker 1977, Cannon et al. 1994) (Augspurger & Franson 1988, Loiselle et al. & equal ground cover Post-logging is different than o flli (Peters 1996) 4 higher colonization levels from (Indonesia, 5 yrs post- ostg seed for wind-dispersed spp. selective: Hill et al. 1995) > Increased dominance of fast growing timber producing tree spp. after silvicultural refinement (Malaysia, MUS: Chua 1998) Reduced leaf biomass but > Vine spp. richness and densities recover to pre-MUS logging and - Reduced dispersal of seeds carried by increased leaf quality; treatment by 40 yrs (Malaysia: Gardette 1998) large-bodied, gap-shy frugivores (Gorchov reduced log distribution, Lich d ir d mo d d 40 d et al. 1993, Forget & Sabatier 1997) or that regenerating stems, and siMcultural treatirent (Maoas div MUS: Wolseley et al0 1998) are scatterhoarded by caviomorph rodents overall trsieeutualteamnt(Mlysa,MSsWize e l.I(Forget & Milleron 199 I; Forget 1990, (Madagascar, RIL, <10% > Palmspp.richnesslowerinloggedandMUStreatedforestthanin 1°forest 1993, 1994; Asquithet al. 1999) esp. if forest: Ganzhorn 1995, (Malaysia Nur Supardi et al. 1998) hunting present Ganzhorn et al. 1990) Vegetation > Fewerlargediametervines40 > Mortality rates 4x higher for spp w/ dbh > yrs post MUS than in II forest 40 cm (S. India, 10-I5 yrs post- selective (Malaysia: Gardette 1998) logging: Pelissier et al. 1998) > Denning trees less available for > Tree mortality increased after arboreal rodents & marsupols liberation/refinement treatments (Costa after selective loggng Rica: Finegan & Camacho 1999) (Auistralia: Laurance & (autra . A > Vine cutting promotes tree growth & Laurance 1996; In vchoa & Soriano in ) reduces logging damage but removes important intercrown pathways for wildlife > Repeated selective logging (reviewed in Putz 1991) reduces abundance of large trees and shifts composition to earlier successional species (Australia: Home & Hickey 1991 Primates > Increased infant mortality > Omnivore densities (1-25 trees harvested/ha, I-SO yrs post-logging, > Herbivory & frugivory - enhanced by low for various spp. Africa, Asia, Amazon) - 8 spp decreased, 13 spp increased, 5 spp intensity (< 10% affected) logging immediately after logging, showed no change; several spp showed opposite results in different disturbance (Madagascar: Ganzhorn 1995); but some spp recovered 6- studies and/or different sites (reviewed in Plumptre & Grieser Johns in data lacking for higher intensities, but 12 yrs post logging press) presumed as variable as changes in spp (Malaysia: Johns & Johns > Folivore/frugivore densities (as above) - 6 spp decreased, 5 spp densities 1995) increased, 3 spp showed no change; as above, several spp showed > Seed dispersal & predation - no specific opposite responses in different studies/sites (Plumptre & Grieser Johns data; probably varies with animal densities in press); hunting affects folivores more than logging itself (Africa: Oates > Vine cutting and liberation thinning eta]. 1996) increased locomotory costs and > Frugivore/folivore densities (as above) - 3 spp decreased, 2 increased, 4 susceptibility to predators for non-volant spp showed no change; again, several spp responded differently in arboreal animals (reviewed in Putz et al. different sites/studies (Uganda: Plumptre & Grieser Johns in press) 2000) > Lemurs in dry forest (< 10% affected by logging) - increased sightings of > Predation of small primates by raptors may all spp attributed to increased fruit & higher foliage quality on remaining increase in response to vine removal trees (Madagascar: Ganzhorn 1995) (Suriname: S. Boinski, pers comm.) > Monkeys in Zaire - 4 spp preferred secondary forest: 3 others preferred primary forest (Zaire: Thomas 199 1) > Folivores - hunting has greater negative impact than logging (Africa: Oates et al. 1996) vp~ > Primate densities decreased in response to logging & MUS (Malaysia: CA1 Laidlaw 1998) Community Attributes Category Structure Composition Function Non-rodent > Browsers/grazers (e.g., Asian elephant, African buffalo, gaur, banteng, & > If no hunting, increased herbivory; if mcmnmds sambar deer) increase (if no hunting) b/c of increased food quality hunting decreased herbivory (Oates et al. 1990, Waterman & Kool 1994) & quantity Johns 1992); the only sensitive spp in this guild was the giant forest hog (Hylochoerus meinertzhageni) (reviewed in Davies et al. in press) > Carnivores - mixed responses & few data; spp with restricted diets most sensitive (reviewed in Davies et al. in press) > Species favored by hunters (e.g., duikers, pigs, peccaries), for special trade (e.g., elephants) and roadside feeders (e.g., Synceros caffer, Bos spp) decline (Davies et al. in press) > Large, slow (moving & reproducing) species decline (hunting a confounding factor) (Lahm 1994, Grieser johns 1997) > Species dependent on fruits/seeds of timber species (e.g. Sus barbatus, Caldecott 1991) on closed canopy (e.g., Muntiocus atherodes, Heydon 1994; Giao et al. 1998), or with limited dietary flexibility due to energetic requirements (e.g., Tragulus spp. Heydon & Bulloh 1997) decline. Rodents and > Arboreal marsupials - I sp declined & 4 spp unaffected after 8- 10 trees (50-55 > Increased seed predation rates (esp if Marsupials m3) harvested/ha (Australia: Laurance & Laurance 1996) hunting) (e.g., Terborgh & Wright 1994) > Small mammal populations most sensitive - according to models: (a) > Decreased predation on large seeds if large Sciuridae, Echimyidae & other canopy dependent frugivores/ granivores; seed predators extirpated (Putz et al. and (b) spp w/ low RA and restricted geog range (Ochoa et al. 1993, 1990) 1995; Utrera 1996; Ochoa 1997; review in Ochoa & Soriano in press) ' Increase in Muridae may increase colonization of small seeded tree spp. Small mammal populations least sensitive - according to models: Didephidae & (Hammond & Thomas in press) Muridae; semi-arboreal omnivores/predators increase (e.g., Didelphis spp. & Philander opossum; see above refs) > Rodent spp richness increased soon after logging (Uganda, 9 trees/ha, 62% canopy open: lsabirye-Basuta & Kasenene 1987, Muganga 1989), but decreased by 16 yrs post (Lwanga 1994) > Rodent spp. richness increased, 2 dominant spp. decreased in abundance but increased in body mass after logging (China: Wu et al. 1996) Bats > Pioneer plant dispersers (e.g., Carollia. Artibeus & Sturnira) increase > Increased dispersal of pioneer plant spp; (Charles-Dominique 1986, Fleming 1988, Fenton et al. 1992, Kikkawa & decreased predation Dwyer 1992, Brosset et al. 1996, Utrera 1996, review in Soriano & Ochoa in press) > Phyllostominae (insectivores & vert predators) - decline (e.g., Artibeus > Reductions in insectivore populations could obscurus, Vampyressa bidens, Phyllostomus elongatus, Tonatia result in increased herbivory rates with sauropila) or disappear (e.g., Vampyrum spectrum, Tonatia sylvicola, deleterious effects on forest resource Chrotopterus auritus, & Peropterix koppleri, Emballonuridae) (Fenton productivity (Soriano & Ochoa in press) et al. 1992, Brosset et al. 1996, Utrera 1996, review in Soriano & Ochoa in press) > Reduced guild complexity & kcal extinctions after logging (5.8 m3/ha, trees > 40 cm dbh); effects exacerbated by enrichment strips (French Guiana: Brosset et al. 1996; similar findirngs in Venezuela: Ochoa in press; reviewed in Soriano & Ochoa in press) > Open-forest, fast-flying spp w/ long, narrow wings do best b/c of increased gap area; dominated 6-yr post site w/ 6-7 trees/ha harvested (Australia: Crome & Richards 1988) > Spp preferring medium to dense understory disappeared after brushing indicating high sensitivity to changes in forest structure (Miller in press) Birds > Nectarivore & generalist frugivore/insectivores - nc or increase in most ) Herbivory & frugivory - reduced for fig- spp (ohns 1989a, 1992b; Lambert 1991; Thiollay 1992; Zakaria 1994, eating spp b/c of lower post-harvest 1996; Mason 1996; reviews in Mason & Thiollay in press, Zakaria & density of Ficus spp (Lambert 1991) Francis in press, & Plumptre in press) > Terrestrial & sallying insectivores (e.g., Tyrannidae) & other spp > Predation - expected to decrease b/c of depending on forest interior (e.g., those in mixed-spp flocks) decrease reduced insect prey abundance (b/c of w/ logging (ohns 1989a, 1991 a, 1992a; Lambert et al. 1992; Thiollay hotter/drier conditions) (Mason 1995) 1992; Zakaria 1994; Bennet & Dahaban 1995; Fanshawe 1995; Grieser Johns 1996; Mason 1996; Owiunji & Plumptre 1998; reviews in Mason & Thiollay in press, Zakaria & Francis in press, & Plumptre in press) Birds > Predators, foliage gleaners & large frugivores - inconsistent responses in > Seed dispersal - reduced b/c of avoidance different sites/studies aohns 1991 a; Lambert et al. 1992; Thiollay 1992; of large openings in forest canopy (Stouffer Grieserjohns 1996; Zakaria & Nordin 1998; reviews in Mason & & Bierregaard 1995a,b) Thiollay in press, Zakaria & Francis in press, & Plumptre in press) > Understory spp. - nectarivorous increased; insectivorous decreased; > Colonization rates -expected to increase latter trend worsened by enrichment planting (Venezuela: Mason 1996) when refuges provided w/in logged areas -(ohns 1996) _____________________________ Community Attributes Category Structure Composition Function > Interior understory spp declined by 37-98%; generalists & spp associated w/ dense second growth, edges & large gaps increased; overall richness and abundance down 27-34% (French Guiana, 1- 17 yrs post, 38% undergrowth & 63% canopy loss: Thiollay 1997); (other study: 3 trees/ha, 10 yrs post) - lower diversity b/c of more uniform habitat (Thiollay 1992) > Flycatchers, wren-warblers, trogons & woodpeckers declined; nectarivores & some frugivores increased (Malaysia, 8 yrs post-logging: Lambert et al. 1992) 9 Spp. richness lower than unlogged (Indonesia: Marsden 1998) > 1/3 spp expected to disappear (Liberia, logged fragment: Kofron & Chapman 1995) Herps > Heliothermic lizards - increased density of large lizards & altered comm > Competition & Predation - increases in structure (Amazonian treefall gaps: Vitt et al. 1998) generalist spp (Vitt et al. 1998) > No change in reptile community comp or abundance (< 10% damage, 2- > Colonization - generalist spp (e.g, Hyla 12 yrs post-logging: Bloxam et al. 1996) geographica) increase (Caldwell 1989) > Lower amphibian diversity (Amazonia, high intensity logging or logging & fragmentation & roads: Pearman 1997, Vitt et al. 1998) > Spp that live in leaf litter most severely affected especially after high intensity logging (e.g., monocyclic harvests); w/ less intense logging most spp persist in short term, but invasion of 2° forest by large predatory lizards may extirpate small frogs & lizards; some frog spp increase after logging b/c of ponding in skid trail ruts or in stream impoundments _._____________ (reviewed in Vitt & Caldwell in press) Insects > Community comp of Araneae, Hymenoptera, Heteroptera, Homoptera, > Short generation times & high fecundity Coleoptera, Orthoptera & Lepidoptera differed in 1° forest, logged allow rapid recovery of effected forest & plantations (Uganda: Nummelin 1996) populations (temperate zone studies: Huhta 1976, Heliovaara & Vaisanen 1984; Spp. restricted to understory, leaf litter & soil most vulnerable (reviewed tropics: Ghazoul & Hill in press) in Ghazoul & Hill in press) Geometrid moth diversity lowest in abandoned clearcut, highest in > Negative cascading effects on insect- forest logged w/ MUS (Malaysia: Intachat et al. 1997; Ghazoul & Hill in decomposition, nutrient cycling) are likely Press) (Ghazoul & Hill in press) > Lower lepidopteran moth diversity after logging + conversion to plantation; Coleopteran dung & carrion beetles less affected (Malaysia: Holloway et al. 1992; Ghazoul & Hill in press) > Dung beetle spp. composition in RIL area resembles 10 forest more than conventionally logged area (Malaysia: Davies in press) > Butterflies, flying beetles, canopy beetles, canopy ants, leaf eater ants, termites and soil nematodes all declined along disturbance gradient (Lawton et al. 1998) > Butterfly richness, abundance & evenness all decreased (Indonesia, 5 yrs post-logging: Hill et al. 1995) > Cassidinae beetle richness greater in logged forest and plantations vs. 1° forest and logged forest community compposition more like plantations (Uganda: Nummelin & Borowiec 199 1) > Opportunistic species increase after invasion of pioneer plants at expense of closed forest spp (Holloway et al. 1992) > Ant spp. richness similar in Io forest and MUS forest 40 yrs after treatment (Malaysia: Bolton 1998) Community Attributes Category Structure Composition Function Soil organisms > Fruiting body (sporome) ) Mycorrhizal fungi spp. richness higher in MUS treated area than in 1° density of mycorrhizal fungi forest (Malaysia: Watling et al. 1998); wood-rotting organisms may be higher 40 yrs. After MUS better impact indicators for logging & other silvicultural treatments treatment than in I' forest > Changes to soil properties directly influences soil fauna diversity (review (Malaysia: Watling et al. in Camilo & Zou in press) 1998) > Endogeic & anecic earthworm communities little affected by small scale clearcuts & low frequency (>50 yr intervals) large scale (reviewed in Camilo & Zou in press). Aquatic systems > Loss of instream & riparian zone diversity; decreased abundance of > Alterations in light & temperature regimes, fishes & benthic invertebrates intolerant of (a) high sedimentation, (b) stream sediment loads, turbidity, stream shifts in photosynthesis/respiration ratios, (c) invasive spp (review in chemistry, hydrology & fluvial Pringle & Benstead in press) geomorphology may cause cascading effects on aquatic food webs (Pringle & Benstead in press) Appendix IV- Bird Responses (densitybased on inter-site comparisons) to Logging in Tropical Forests, Organized by Feeding Guild Biodiversity Series - Impact Studies 51 01 Bird responses (density' based on inter-site comparisons) to logging in tropical forests, organized by feeding guild Ulu Ulu Budongo, Kibale, Tekam, Segama, Segama, Imatoca, Piste de St -Elie, Site Uganda Uganda Malaysia Sabah Sabah Venezuela French Guiana Logging intensity 20-80 m3/ha 2! m3lha 18 trees/ha 79 m3lha 90 m3lha 5-8 m3lha 1O m3lha 10 m3/ha Recovery (years since harvest) 25-50 25 6-7 3-4 9-10 1-6 1 10 Hunting no no unknown no no no yes yes Guild Frugivore + nc + - nc nc - - Frugivore/insectivore (arboreal) nc + + + + + nc nc Frugivore/insectivore (terrestrial) nc + - - nc nc na na Frugivore/granivore nc nc na na na na na na Granivore nc + na na na + na na Granivore/insectivore nc + na na na na na na Insectivore (sallying) - - - nc nc Insectivore (terrestrial) nc nc Insectivore (arboreal foliage gleaner) nc nc - nc na Insectivore (understory foliage gleaner) nc + nc - nc Insectivore (bark gleaning) nc - - nc Insectivore/nectarivore nc nc nc + + + Predator nc + nc + - na Predator/frugivore na na - nc nc na Source Owiunji and Dranzoa Johns 1989a Nordin and Lambert Mason 1996 Thiollay Thiollay Plumptre 1995 Zakaria et al. 1992 1992, 1997 1992,1997 1998 1997 Notes: Density (calculated by same method within a study, but differed between studies). + = higher following logging, - = lower following logging, nc = no change, na = information not available; P B 0.05. (Adapted from Mason and Thiollay, in press; Plumptre et al., in press; and Zakaria and Francis, in press). Appendix V Impacts of Logging (and other silviculture treatments where noted) on the Species Component of Biodiversity in Tropical Forests Biodiversity Series - Impact Studies 53 Impacts of logging (and other silviculture treatments where noted) of the species component of biodiversity in tropical forests Species Attributes Taxa Structure Composition Function Vegetation > Smaller size classes most affected > High grading large commercial trees > Reduced seed availability & reduced seed dispersal by rodents of by felling, skidding, and road caused local extirpation of Caesalpinia large seeded commercial species after logging (Guyana: Forget & construction echincata, Ceiba pentandra, and Aniba Hammond in press, Hammond & Thomas in press) > 62% seedling & 59% sapling duckei; Amazonia (Gentry & Vasquez > For dioecious species, seed crop decreases with increasing mortality (Malaysia: Borhan et al. 1988, Grieser Johns 1997) distance of males to females (Mack 1997) 1987) > Harvesting figs promotes their ) Higher seedling mortality (esp. in very disturbed areas) in Sabah S I 1I% of trees uprooted (Brazil: regeneration & may not have wildlife (Pinard et al. 1996) Uhl & Guimaraes 1989) impacts due to scarcity of well-formed (iade l 96 trees (Bolivia; Fredericksen et al. 1999) > Lower seedling growth rates (Nussbaum et al. 1995) 17 trees damaged for every tree > Populations of wild fruit trees are often ) Reduced density of adults (i.e. breeding individuals) 4 fewer cut (E. Kalimantan: Abdulhadi et decimated by destructive harvesting (Peru: flowers per unit area (C. Peters, pers. comm.) al. 1981) Vazquez & Gentry 1989) > 1% increase in tree mortality 3-4 yr post-logging (Silva et al. 1995, 0 >50% of young trees killed (N. F1 & Camacho 1999) Queensland, Australia: Crome et Fegan al. 1992) ) 4-fold increase in tree mortality 2 yrs post-logging (Thiollay 1992) )0- - 60-70% of residual trees ) More resources available to residual trees b/c of reduction in damaged by logging (Fox 1968; canopy cover and stem density (Maitre 1991); but damaged trees Nicholson 1979) may need to use resources to heal w/ net result of increase in > Large and small trees killed with aborted fruits (Stephenson 1981) equal probability Johns 1988; > Potentially higher seed predation levels (Schupp 1990, C. Peters, Crome et al. 1992) pers. comm.) > Species w/ type IIl population > Temporary increase in growth rates depending on extent of structure may be easily canopy opening (Wan Razali 1989) eliminated (C. Peters pers. > Mean dbh increment 2-3 x higher in logged vs. unlogged Jonkers comm.) 1987) > Spatially clumped pattern of > Lower growth rates after thinning + selective logging in (E. conspecifics shifts to dispersed or Malaysia: Primack et al. 1989) random patterns 4 reproductive biology and logistics of future > Lower growth rate for Araucaria cunninghmamii (Australia: Enright silviculture 1978) Primates > > See Appendix VI Non-rodent > Civet densities in logged forest were only mammals 20% of 1° (Malaysia, 2-12 yrs post selective logging, no hunting: Heydon & Bulloh 1996); but increased when <33- 50% forest impacted (Malaysia, 5- 10 yrs post logging: Johns 1983b) > Tree shrew densitites greater 40 yrs post MUS treatment than 1 0 forest (Malaysia: Laidlaw 1998) > Giant forest hog populations decline after logging (Davies et a]. in press) > Elephant populations increase in response to logging (Africa & Asia: Grieser Johns 1997) > Mousedeer (Tragulus) populations reduced after logging (80- 100 m3/ha, Malaysia: Heydon & Bulloh 1997) Rodents & > Squirrel & rat populations showed variable Marsupials & species-specific responses to MUS (Malaysia, 40 yrs post-treatment: Laidlaw 1998) Bats > Vampire bat densities increase if cattle introduced to logged areas aohns et al. 1985, Wilkinson 1985, Johns 1988, 1992a, Fenton et al. 1992) Birds > See Appendix IV Herps Insects Soil organisms Appendix VI Primate Responses (by feeding guild) to Logging in Tropical Forests Biodiversity Series - Impact Studies 57 Primate responses (by feeding guild) to logging in tropical forests; O=no change; - = decrease after logging; + = increase after logging GUILD / SPECIES Ulu Segama Nanga Gaat Tekam Sg. Lalang Kemasul Pta da Kibabe Budongo Kalinzu Kirindy Lope Castanha Logging Intensity (trees/ha) 20 10 18 > 20 >20 5.1 or 7.4 6-25 ? . I Damage levels (% tree loss) 32-58 54 5 38 55 60 25 or 50% >50 ? II Years since logging 6-12 0-4 0-12 20-30 20-30 11 12-28 1-50 4-20 1 -15 Omnivores Cebus apella + Cebus albifrons + Lophocebus albigena 0/- 0 Cercopithecus Ihoesti O_0 Cercopithecus ascanius +1- + + I Cercopithecus cephus 0 Cercopithecus medius + Cercopithecus mitus _ 0/- + 0 Cercopithecus pogonias 0 Cercopithecus nictitans . 0 Ga/ago s_p . Gorilla gorila 0 Macaca spp _ Macaca SPP _ Macaco fascicularis 0 + . MacOca nemestrina 0 0 + 0 Mandrillus sphinx 0 Microcebus spP + Pan trogtodytes 0/- 0 + + Perodicticus potto . Phaner furcifer _ + Pongo pygmaeus + Saguinus mystax + Saimiri sPP _ + Folivores/frugivores Alouatto seniculus _ Ateles paniscus 0 Callicebus moloch + Callicebus torguatus __ __ + Procolobus badius 0/- Colobus guereza . + + 0 Colobus satanus 0 Presbytis hosei + . Presbytis melalophos + _ Presbytis obscura + Presbytis rubicunda . . Frugivores/folivores I I l_l Hylobates tars + Hylobates muelleri 0 _ Lagothrix lagotricha Pithecia albicans + _ References Johns Dahaban Johns Laidlaw 1994 Laidlaw 1994 Johns 1991 b Skorupa 1986, Plumptre & Howard Ganzhorn White 1992, 1989a,b 1996 1989a,b 1988; Weisenrseel Reynolds 1991; 1995; 1994 et al. 1993; 1994 Hashimoto Ganzhorn et Chapman et al. 1995 al. 1990 * Modified from Plumptre and Grieserjohns (in press) u- Appendix VII Impacts of Logging (and other silviculture treatments where noted) on the Genetic Component of Biodiversity in Tropical Forests Biodiversity Series - Impact Stutdies 61 Impacts of logging (and other silviculture treatments where noted) on the genetic component of biodiversity in tropical forests Genetic Attributes Taxa Structure Composition Function Vegetation P Genetic diversity decreased by ' Outcrossing rates unchanged in Shorea leprosula (Chan 198 1, 5.0-23.4% (logged I yr prior; Lee et al. 1996, Wickneswari et al. 1997) & Dryobalanops regenerated stands showed no aromatica (Kitamura et al. 1994) significant loss); effect on dioecious Outcrossing rates decreased in Shorea megistophylla (Murawski species perhaps underestimated et al. 1994a,b); density of mature individuals likely an important (Wickneswari et al. 1997) factor > Reduced cross-pollination rates in Shorea siamensis after logging (Ghazoul et al. 1998) and reduced pollination success of Dipterocarpus obtusifolius after logging S. siamensis (Ghazoul 1999) P Outcrossing rates of Carapa spp. were lower after controlled experiments in logged vs. unlogged forests in French Guiana (Doligez & Joly 1997) but were not significantly different in Carapa stands in Costa Rica (Hall et al. 1994) possibly because of very high stand densities in Costa Rica. P Studies citing genetic erosion of Swietenia mahoganii from logging have not been substantiated in follow-up studies (reviewed in Lugo 1999) Primates Non-rodent mammals Rodents & Marsupials Bats Birds Herps Insects Soil organisms _s_ _ Glossary' Advanced regeneration: seedlings or saplings compound) interest rate; used to calculate the present in the understory prior to logging or "net present value" other silvicultural treatment Edaphic: related to or caused by particular Aerial logging: a timber yarding system using conditions in the soil suspended cables, helicopters, or balloons to lift logs Enrichment planting: increasing the density of desirable species by interplanting, planting in Afforestation: establishment of forests in areas gaps, or planting along cleared lines that did not historically support them (e.g., in savannas or drained marshes) Fragmentation: the process of breaking once continuous expanses of an ecosystem type Bole: trunk or main stem of a tree (typically forest) into island-like patches in a matrix of other land uses Buffer zone: vegetation maintained on the borders of roads, wetlands, or other ecologically Gap: generally refers to a hole in the canopy sensitive areas to mitigate the impacts of created by the death of a canopy tree, but there management activities are also understory gaps and belowground gaps Bulk density: the weight per unit volume of a Girdle: to make an incision around a tree trunk material (often soil) at least as deep as the cambium with the intention of killing the tree; if combined with Carbon sequestration: carbon storage arboricide application then referred to as "poison girdling" Coppice: sprouting of trees from stumps or roots Group selection: harvesting of trees in clusters generally no wider than twice the height of the Cutting cycle: the time (years) between stand mature trees to promote regeneration of entries for timber extraction; there can be more moderately light-demanding species (a than one cutting cycle in a "rotation" "polycylic" system) Dendrology: the study of trees including their Hauling: carrying logs by truck (lorrie) from the identification forest to processing or exporting facilities Discounting: adjusting a future value by Heartrot: decomposition of the central dividing by a function of the (generally stemwood of living trees typically due to fungal Biodiversity Series - Impact Studies 63 Biodiversity Conservation in the Context of Tropical Forest Management infection after mechanical or fire-related Reduced-impact logging: a set of techniques damage designed to avoid excessive damage to soil or to the residual stand during and after timber Heterozygous: having two different alleles at harvesting; typical components include pre- the same locus on homologous chromosomes; planned skid trails, directional felling, and an index of genetic diversity engineering specifications for stream crossings High-grading: removal of the best trees often Reforestation: the recovery of forests in areas leaving a residual stand dominated by trees of that were deforested; compare to "afforestation" poor form and non-commercial species; "creaming" Regeneration method: a timber harvesting method that promotes development of a new Highlead yarding: a yarding system in which age class; the principal methods include logs are skidded along the ground to a central clearcutting, seed tree, shelterwood, selection, point by a cable passing through a block at the and coppicing top of a tower or spar tree (not to be confused with "skyline yarding") Riparian: a terrestrial area bordering a water body Liana: woody vine Rotation: the period between establishment of regeneration and final harvesting in even-aged Liberation thinning: a silvicultural treatment in stands which future crop trees are released from competition typically by girdling near neighbors Shelterwood method: a stand regeneration (note: term used differently by many North method that occurs in two phases, a first phase American foresters) in which enough timber is harvested to promote development of a new age class in partial shade, Logging intensity: can refer to either the timber and a final felling of the remaining mature volume or number of trees harvested per unit timber after regeneration is established area Silviculture: the art and science of controlling Natural forest management: management of the establishment, growth, composition, and forests for timber, non-timber forest products, health of forests and woodlands and environmental services by relying principally on natural regeneration (compare to Skidding: yarding logs with a rubber-tired self- plantation forestry) propelled machine or bulldozer ("snigging" in Australia) Neotropics: American tropics Skyline yarding: a log yarding system in which Net present value: the current value of some logs are partially or entirely suspended from a future cost or benefit taut cable (not to be confused with "high-lead yarding") Paleotropics: tropical areas in Africa, Asia, Australia, and the Pacific Islands Stand: a contiguous group of trees with a more- or-less similar history of disturbance, species Plantation: a stand of trees established by composition, size-class distribution, and planting; 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